#ZooKeys
ZooKeys 1201: 167-217 (2024) DOI: 10.3897/zookeys.1201.120750
Research Article
An island in a sea of sand: a first checklist of the herpetofauna of the Serra da Neve inselberg, southwestern Angola
Mariana P. Marques'2*4®, Diogo Parrinha22“®, Manuel Lopes-Lima2*®, Arthur Tiutenko®®, Aaron M. Bauer®®, Luis M. P. Ceriaco2?”®
1 Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh PA 15213, USA 2 CIBIO, Centro de Investigagao em Biodiversidade e Recursos Genéticos, InBIO Laboratorio Associado, Campus de Vairao, Universidade do Porto, 4485-661
Vairado, Portugal
N DOD oo SP W
BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairado, 4485-661, Vairao, Portugal
Departamento de Biologia, Faculdade de Ciéncias da Universidade do Porto, Rua do Campo Alegre 1021, 4169-007 Porto, Portugal Friedrich-Alexander-Universitat Erlangen-Nurnberg, Schlossplatz 4, 91054 Erlangen, Germany
Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista, Sao Cristdvao, 20940-040 Rio de Janeiro, RJ, Brazil
Corresponding author: Mariana P. Marques (mptimarques@gmail.com)
OPEN Qaccess
Academic editor: Anthony Herrel Received: 13 February 2024 Accepted: 11 March 2024 Published: 14 May 2024
ZooBank: https://zoobank. org/23C7E6E7-AE73-4685-AEDA- 26DEBOEE0068
Citation: Marques MP Parrinha D, Lopes-Lima M, Tiutenko A, Bauer AM, Cerfaco LMP (2024) An island in a sea of sand: a first checklist of the herpetofauna of the Serra da Neve inselberg, southwestern Angola. ZooKeys 1201: 167-217. https://doi. org/10.3897/zookeys.1201.120750
Copyright: © Mariana P. Marques et al. This is an open access article distributed under terms of the Creative Commons Attribution
License (Attribution 4.0 International - CC BY 4.0).
Abstract
The Serra da Neve inselberg in Namibe Province, southwestern Angola is the second highest peak of Angola with an elevation of 2489 m. It remains one of the least explored regions in the country, despite several endemic species having been recently described from this inselberg. Here we provide an inventory of the amphibian and reptile species ocurring in Serra da Neve and compare its fauna with that of the surrounding habitats at lower elevations. We also examine the phylogenetic affinities of the inselberg taxa. A total of 59 herpetological taxa were recorded for the Serra da Neve inselberg and its immediate surroundings. These include 11 species of amphibians, belonging to nine genera and seven different families, and 48 species of reptiles, belonging to 32 genera and 12 families. Of these, one amphibian and seven reptiles from seven different genera are strictly endemic, making the inselberg the richest region in southwestern Africa with respect to strict endemics, with one endemic reptile taxa per 127 km?. Not surprisingly, most of the recorded taxa belong to clades that are endemic, or at least strongly asso- ciated, with southern Africa, but two are representatives of central African clades, and another two are more closely related to eastern African highland taxa. We also provide comments on the threats to the conservation of this endemic-rich inselberg.
Key words: Amphibians, Angola, biodiversity, conservation, endemism, inselberg, reptiles, Southwestern Africa, taxonomy
Introduction
Inselbergs are isolated mountains/rock outcrops which rise more or less abruptly above a plain. Scattered across all continents, these rock outcrops are usually im- portant biodiversity hotspots, serving as refugia for diverse plant and animal taxa (Porembski and Barthlott 2000; Burke 2005; Brand et al. 2011; Bayliss et al. 2014; Porembski et al. 2017). Acting as islands, inselbergs are usually separated and
167
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
isolated from each other in a way similar to oceanic islands (MacArthur and Wil- son 2001). Due to their isolation and geomorphologic delimitation from the sur- roundings, for some groups they represent partly independent ecosystems that are especially suitable for comparative research on the structure and dynamics of floral and faunal communities (Porembski and Barthlott 2000). Their microcli- matic conditions and habitats, which usually differ from those in the surrounding lowlands, allow inselbergs to support unique biological communities and high levels of endemism (Simmons et al. 1998; Porembski and Barthlott 2000; Burke 2001, 2003, 2005; Porembski 2007; Michael et al. 2008; Brand et al. 2011, 2019; Bayliss et al. 2014). In addition, they are excellent models for studying aspects of island ecology, geographical differentiation and mechanisms influencing diversi- ty. Due to their topography, Inselbergs are typically not used for agriculture and, except for some instances of tourism, have not been transformed by human ac- tivities. They thus constitute almost pristine habitats with special importance for conservation, and usually provide favorable conditions for flora and fauna, con- cerning availability of water, shade, and refuge (Porembski and Barthlott 2000).
The study of African inselbergs and other “sky-islands” has been a topic of great interest in recent years for a broad scope of scientists, from conserva- tion biologists to systematists. This is especially true for East-African Afrom- ontane inselberg regions, as in the case of Mount Namuli and Mount Mabu in Mozambique (e.g., Timberlake et al. 2009; Portik et al. 2013a, 2013b; Bayliss 2014) or Mount Mulanje in Malawi (Curran et al. 2012). Due to this interest, in recent years several new endemic animal species, such as amphibians (Ceria- co et al. 2018; Conradie et al. 2018), reptiles (Branch et al. 2005; Branch and Bayliss 2009; Branch and Tolley 2010; Portik et al. 2013b; Branch et al. 2014, 2019; Marques et al. 2019, 2020, 2023a, 2023b), invertebrates (Congdon et al. 2010; Daniels and Bayliss 2012; Bilton 2014; Daniels et al. 2014, 2020), and mammals (Monadjem et al. 2010; Taylor et al. 2012; Simmons et al. 2021) have been described from inselbergs across Africa. Several studies have also pro- vided inventories of the herpetofauna (Michael et al. 2008; Kirchhof et al. 2010; Conradie et al. 2016a; Bittencourt-Silva et al. 2020) and plant diversity (Rabari- manarivo et al. 2019; Porembski and Barthlott 2000; Burke 2003; Kandziora et al. 2022; Brand et al. 2019) of these inselbergs across the continent.
In contrast to East-African Afromontane inselberg studies, xeric southwest- ern African inselbergs have been largely neglected in the fauna studies (Elzen 1983; Griffin 2000), with most focusing only on flora and vegetation (Porembski and Barthlott 2000), and, more recently, on carnivores (Rapson et al. 2013). Northwestern Namibia and west-central Angola form an area with a consid- erable diversity and number of inselbergs. Surrounded by desert and savanna habitats, southwestern African inselbergs are usually rocky outcrops of diverse geology and origins that contrast with the intervening habitats (Goudie and Viles 2015). Both reptile diversity and endemism are high in rocky areas in An- gola and Namibia (Bauer et al. 2023).
The landscape of southwestern Angola is characterized by the presence of isolated mountain-like rocky outcrops of subvolcanic origin mostly composed of gneisses, migmatites and granites (Pereira 1977). One of the most impressive southwestern Angolan inselbergs is Serra da Neve. Located at the northern limit of Namibe Province, southwestern Angola, and with a basal area of approximately 630 km’, Serra da Neve is the second highest peak of Angola, with an elevation of
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 168
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
2489 m (Pereira 1977). It lies in what Mendelsohn and Huntley (2023) define as “the southern escarpment landscape”, an area ranging from the Coporolo River in Ben- guela Province, Angola, to the Huab River in Namibia. The inselberg is covered by a Miombo forest habitat, contrasting with the surrounding lowland habitats, which are mainly dominated by Namibian woodland savanna and arid areas of Namib Desert (Grandvaux-Barbosa 1970). Although its biodiversity is still poorly known, and systematic surveys have only recently begun, the inselberg is already known to harbor an impressive number of strictly endemic species of amphibians and reptiles, such as Poyntonophrynus pachnodes Ceriaco, Marques, Bandeira, Agar- wal, Stanley, Heinicke, Blackburn & Bauer, 2018, Cordylus phonolithos Marques, Ceriaco, Stanley, Bandeira, Agarwal & Bauer, 2019, Lygodactylus baptistai Marques, Ceriaco, Buehler, Bandeira, Janota & Bauer, 2020, Afroedura praedicta Branch, Schmitz, Lob6n-Rovira, Baptista, Antonio & Conradie, 2021, and Acontias mukwan- do Marques, Parrinha, Tiutenko, Lopes-Lima, Bauer & Ceriaco, 2023 (Ceriaco et al. 2018; Marques et al. 2019, 2020, 2023b; Branch et al. 2021), with at least three oth- er reptile species currently being described (MPM unpubl. data; DP unpubl. data). Currently, no data for other taxonomic groups exist, but recent multidis- ciplinary surveys have also uncovered interesting and cryptic diversity within those, which will result in a better understanding of the taxonomic diversity, biogeographic patterns, and endemism of Serra da Neve. In this context, the main objective of this study is to provide a first description of Serra da Neve her- petofauna, and more specifically to 1) to provide an inventory of the occurring species as well as list taxa that have not yet been recorded, but which may be present; 2) to compare the fauna present on the inselberg with that of the sur- rounding lower elevation habitats; 3) to examine the phylogeographic affinities of the inselberg taxa; 4) to compare the level of endemism of Serra da Neve with other regions in Angola and southern Africa; and 5) to present a first glimpse into the major conservation threats that the inselberg herpetofauna may be facing.
Materials and methods
Three herpetological surveys of Serra da Neve and its surrounding areas have been carried out since 2016. The first survey was conducted from 18 to 22 No- vember 2016, the second one from 26 to 28 February 2019, and the third one from 26 October to 6 November 2022. A total of eight main sites were surveyed (Table 1, Fig. 1). The combination of sites was chosen to maximize the types
Table 1. Sampling localities in and around the Serra da Neve inselberg, and respective latitude, longitude, and elevation data.
Localities Latitude, Longitude Elevation (m) Map (see Fig. 1) Road to Quilengues -13.8159, 13.3264 587 1 N’Dolondolo 13-8133; 43;1362 681 2 Mamué -13.8003, 13.1229 701 3 Maylowe -13.6355,.13:2755 798 4 2 km N of Maylowe -13.8280, 13.2625 820 5 Basecamp 1 =19:7770, 13.2591 1488 6 Lutala Crater -13.7325, 13.1841 1567 ve Catchi -13.7627, 13.2564 1590 8
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 169
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
13.0°E 13.1°E 13.2°E 13.3°E
Elevation (m) ) 250 - 500
(_] 500 - 750
[_] 750 - 1000 [] 1000 - 1250 [) 1250 - 1500 MH) 1500 - 1750 MH 1750 - 2000 | (1 2000 - 2250 L_] 2250 - 2500
DEMOCRATIC REPUBLIC ‘OF THE CONGO
13.0°E 13.1°E 13.2°E 13.3°E
Figure 1. Map of the sampling localities in Serra da Neve and its surroundings; contour lines represent elevation in me- ters above sea level. Localities 1 road to Quilengues 2 N’Dolondolo 3 Mamué 4 Maylowe 5 2 km N of Maylowe 6 base- camp 1 7 Lutala Crater 8 Catchi.
of habitats surveyed, including rocky outcrops, woodlands, open grasslands, streams, and ponds, and to capture different elevations from the base to the top of Serra da Neve. Locality data are presented in decimal degrees using the WGS-84 map datum, and elevation is presented as meters above sea level.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 170
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
The areas surrounding the base of Serra da Neve are all dominated by rel- atively dense Mopane (Colophospermum mopane) woodlands on sandy soil (Fig. 2a—d; Huntley 2023). This habitat dominates the landscape around May- lowe village and extends for a considerable radius around Serra da Neve. The Mopane trees can be seen up to ~ 1100 m elevation on the inselberg, where they are replaced by a largely intact Miombo woodland that dominates the landscape at higher elevations. Still, at the base of Serra da Neve, some areas have interesting vegetation and geological characteristics that are worth men- tioning. The N’Dolondolo area, while still dominated by Mopane woodlands, is considerably more humid than other areas near Maylowe, especially due to its
Figure 2. Lowland habitats in the near surroundings of the Serra da Neve inselberg a Mo- pane habitat in the vicinity of Maylowe b dry Mopane habitat, 2 km N of Maylowe c sandy areas with Mopane leaflitter near Maylowe d dry riverbed near Maylowe e, f N’Dolondolo g, h riparian vegetation in Mamué. Photographs by LMPC (a), AT (b-d) and IA (e-h).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 171
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
ZooKeys 1201: 167-217 (2024), DOI:
Figure 3. Highland habitats in Serra da Neve inselberg a Miombo woodland in the vicinity of Catchi b preserved Miombo woodlands on the way to Lutala c agricultural crops on the way to Lutala d disturbed landscape, vicinity of Catchi e Miombo woodland with gran- ite outcrops, vicinity of Catchi f riparian vegetation on the way to Lutala crater g, h sparse Miombo savanna habitat at Lutala crater. Photographs by LMPC (a-c) and AT (d-h).
sulphurous hot water spring, and its soil is less sandy, with outstanding granite outcrops present (Fig. 2e, f). Further north-west from N’Dolondolo, but still at the base of Serra da Neve, the Mamué area already presents a more complex habitat, with Mopane still present, but the landscape dominated by streams and associated riparian vegetation that extends downward from Serra da Neve, with large waterfalls (Fig. 2g, h).
As elevation rises, the landscape becomes completely dominated by what Grandvaux-Barbosa (1970) described as sparse Miombo woodlands, with the presence of Brachystegia and Julbernardia trees, and the soil becomes rockier
172
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
and with the conspicuous presence of large granite outcrops (Barker et al. 2015). At an elevation of ~ 1490 meters, near Basecamp 1, the area is still well-preserved, without much anthropogenic degradation. There is almost no grass, but different types of bushes (Combretum spp.) are present and there is a considerable accumulation of leaf-litter below the tree canopy. This type of habitat is continuous throughout the largest part of the inselberg above 900 m (pers. Obs.). The closest areas (~ 100 km) where this type of vegetation occurs are in the margins of the Escarpment around Cubal, Chongoroi and Quilengues, and then further inland in Cangandala, in the Queve and Kwanza River valleys (Grandvaux-Barbosa 1970). Catchi (1590 m), an important collecting locality in these surveys, is a small village and human impacts on the landscape are no- table, as most of the plateau around the village is grazed and transformed into corn and maize plantations, or cattle pastures, with the Miombo woodlands restricted to the steeper slopes around the village. A small stream, with its re- spective riverine gallery, passes through the Catchi plateau, adding to the com- plexity of the landscape. In the main crater of the inselberg near Lutala village (1567 m), tree density is notably lower, forming an open Miombo savanna with herbaceous undergrowth, and fewer shrubs and granite outcrops in contrast to other areas of the mountain (Fig. 3).
Specimens were collected using pit fall traps, long-nooses, rubber bands, or by hand during both diurnal and nocturnal visual encounter surveys. Pitt falls were set in two different sites in a dry riverbed near Maylowe, Serra da Neve base. Each pit fall consisted of a line of four buckets, active for three days (3-5 Nov 2024). All specimens were euthanized following Villanova University animal care and use protocol #1866, preserved in 10% buffered formalin in the field, and then gradually transferred to 70% ethanol for long term storage. Liver tissues were extracted before formalin fixation and preserved in 95% ethanol. Vouch- er specimens were deposited in the herpetological collections of the California Academy of Sciences, USA (CAS); Florida Museum of Natural History, USA (UF); Museu Nacional de Historia Natural e da Ciéncia, Universidade de Lisboa, Portu- gal (MUHNAC/MB), Museu de Historia Natural e da Ciéncia da Universidade do Porto, Portugal (MHNC-UP/REP), and a subset of specimens were deposited in the reference collection of Instituto Nacional da Biodiversidade e Areas de Con- servacao (INBAC) in Luanda, Angola. In some cases, species identifications were further confirmed by sequencing the mitochondrial 16S ribosomal RNA gene.
Results
A total of 59 herpetological taxa were recorded for the Serra da Neve inselberg and its immediate surroundings. These include 11 species of amphibians, be- longing to nine genera and seven different families, and 48 species of reptiles, belonging to 32 genera and 12 families. No crocodilians were recorded in the area. The families Scincidae and Gekkonidae were those represented by the largest number of species (14 and 10, respectively). Among the recorded spe- cies, 22 were found exclusively in the lowland areas of the inselberg base and 14 in the highlands, while 23 species were recorded throughout the study area (Table 2). Eight species are strictly endemic to the Serra da Neve inselberg, and 23 additional species are regional endemics to southwestern Angola and central and northwestern Namibia (Table 2).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 173
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Table 2. Synoptic table listing all recorded species, with notes on elevational distribution and endemicity. Lowland localities in- clude Mamué, Maylowe and its surroundings, N’Dolondolo, and the road to Quilengues (below 1000 m above sea level), while highland localities refer to Catchi, Basecamp 1, and Lutala Crater (> 1000 m above sea level; see Table 1 for further details).
Taxa Serra da Neve | Serra da Neve Strict endemic Regional endemic (southwestern Angola lowlands highlands and central and northwestern Namibia) AMPHIBIA Anura Family Pipidae Genus Xenopus Xenopus petersii X x
Family Bufonidae Genus Poyntonophrynus Poyntonophrynus grandisonae Xx X Poyntonophrynus pachnodes 8 X Genus Sclerophrys Sclerophrys pusilla X x Family Microhylidae Genus Phrynomantis Phrynomantis annectens x Family Arthroleptidae Genus Leptopelis Leptopelis anchietae X X Family Ptychadenidae Genus Ptychadena Ptychadena anchietae X Family Phrynobatrachidae Genus Phrynobatrachus
Phrynobatrachus natalensis X Xx Family Pyxicephalidae Genus Amietia
Amietia angolensis X Genus Tomopterna
Tomopterna ahli x
Tomopterna tuberculosa X X REPTILIA Testudines
Family Testudinidae Genus Kinixys Kinixys belliana X X Genus Stigmochelys Stigmochelys pardalis X Squamata Family Gekkonidae Genus Afroedura Afroedura praedicta Xx x Genus Hemidactylus Hemidactylus benguellensis X Xx X Genus Chondrodactylus Chondrodactylus pulitzerae X X X Genus Lygodactylus Lygodactylus baptistai X x X Lygodactylus nyaneka X X
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 174
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
lowlands highlands Genus Pachydactylus Pachydactylus caraculicus a | |e Pachydactylus maiatoi a a a | | Pachydactylus cf. punctatus _———E_ Genus Rhoptropus Rhoptropus aff. barnardi fx Vit ke si Rhoptropus aff. montanus a a |
Family Lacertidae Genus Heliobolus Heliobolus crawfordi a a aan Genus Pedioplanis Pedioplanis haackei ——— SS EE | Pedioplanis serodioi a |
Family Cordylidae Genus Cordylus Cordylus phonolithos Family Gerrhosauridae Genus Cordylosaurus
Genus Gerrhosaurus
Genus Matobosaurus
Matobosaurus maltzahni i | | Family Scincidae Genus Acontias
Genus Mochlus
Genus Panaspis Panaspis mocamedensis Sl is |i
Genus Sepsina
Genus Trachylepis Trachylepis albopunctata a ie a Trachylepis ansorgii SS eS ae Trachylepis binotata | ie ||
Trachylepis bouri
——
Family Chamaeleonidae
Genus Chamaeleo
Zookeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750
Regional endemic (southwestern Angola and central and northwestern Namibia)
~ | K | mK | OK] CK] OK
175
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Taxa
Family Agamidae Genus Agama Agama aculeata Agama schacki Serpentes Family Typhlopidae Genus Afrotyphlops Afrotyphlops schlegeli petersii Family Leptotyphlopidae Genus Leptotyphlops Leptotyphlops cf. scutifrons Family Pythonidae Genus Python Python natalensis Family Viperidae Genus Bitis (Subgenus Macrocerastes) Bitis (Macrocerastes) gabonica Genus Causus Causus nasalis Family Lamprophiidae Genus Boaedon Boaedon variegatus Genus Hemirhagerrhis Hemirhagerrhis viperina Genus Lycophidion Lycophidion hellmichi Genus Psammophis Psammophis subtaeniatus Psammophylax tritaeniatus Family Colubridae Genus Dasypeltis Dasypeltis scabra Genus Telescopus
Telescopus semiannulatus polystictus
SerradaNeve Serra da Neve
lowlands highlands
X X X X
X X
X X X X X X X X X X
Endemicity levels
Strict endemic
Regional endemic (southwestern Angola and central and northwestern Namibia)
Serra da Neve currently harbors a total of eight strictly endemic herpetologi- cal species (one amphibian and seven reptiles, from seven different genera). We consider strictly endemic taxa to be those currently only known to occur within the area defined by the base of the Serra da Neve inselberg (see Fig. 1). These include one species of bufonid frog Poyntonophrynus pachnodes (family Bufonidae), one cordylid lizard, Cordylus phonolithos (family Cordylidae), three geckos, Lygodactylus baptistai, Afroedura praedicta, and Rhoptropus aff. monta- nus (family Gekkonidae), and three skinks, Acontias mukwando, Panaspis sp. 1,
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750
176
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
and Panaspis sp. 2 (family Scincidae). No strictly endemic snakes or chelonians are known from Serra da Neve. Besides the strictly endemic taxa, the inselberg hosts a number of other regional highland endemics, such as Leptopelis anchi- etae (Bocage, 1873), Trachylepis huilensis (Laurent, 1964), Trachylepis ansorgii (Boulenger, 1907), Trachylepis bouri Ceriaco, Marques, Parrinha, Tiutenko, Wei- nell, Butler & Bauer, 2024, and Pachydactylus maiatoi Marques, Parrinha, Ceria- co, Brennan, Heinicke & Bauer, 2023, all associated with the highland areas of southwestern Angola.
When numbers of strictly endemic taxa on Serra da Neve and the other southwestern African highlands are compared, the inselberg stands amongst the richest in the region (Table 3; Bauer et al. 2023; Becker et al. 2023). Serra da Neve hosts a total of seven strictly endemic reptile species, the highest level of endemicity in the country. The Huila Escarpment and Plateau in southwestern Angola (with an approximate area of 18000 km?), harbor only five strictly en- demic reptile species (Table 3; Bauer et al. 2023; Becker et al. 2023). All other inselbergs, both in Angola and Namibia, have a maximum of two strictly en- demic species (Table 3; Bauer et al. 2023; Becker et al. 2023). Serra da Neve's strictly endemic numbers are even more striking when considering its area. With an approximate area of 630 km?, Serra da Neve is undoubtedly the rich- est region in southwestern Africa with respect to strict endemics per unit of area, with one endemic reptile taxa per 127 km? (Table 3; Bauer et al. 2023), and much more similar to or higher than those found on endemic-rich oceanic islands (Ceriaco et al. 2022; Bauer et al. 2022), or South American table moun- tains, known as “tepuis” (Recoder et al. 2020; Fouquet et al. 2023).
Biogeographic affinities
Not surprisingly, most of the recorded taxa belong to clades endemic to, or at least strongly associated, with southern Africa. The exceptions are Bitis (Mac- rocerastes) gabonica (Duméril, Bibron & Duméril, 1854), Panaspis cabindae (Bo- cage, 1866) Acontias mukwando, and Lygodactylus baptistai, of which the first two are representatives of central African clades, and the latter two are more closely related to eastern African highland taxa. The subgenus Macrocerastes, of which large-bodied vipers such as Bitis nasicornis (Shaw, 1792), B. rhinocer- os (Schlegel, 1855), B. parviocula (BOhme, 1977) and B. gabonica are members, is a group predominantly associated with central African habitats (Barlow et al. 2019; Ceriaco et al. 2020a), even if some of these species have some popula- tions in southern (B. gabonica) and eastern Africa (B. parviocula). Within this group, is B. heraldica (Bocage, 1889), a species endemic to the Angolan central highlands, and the only member of the subgenus which is small-bodied (Ceria- co et al. 2020a). The specimen of B. gabonica from Serra da Neve represents the southwestern-most record of the subgenus in the continent.
A similar distribution pattern can also be observed for Panaspis cabindae and the putative new species Panaspis sp. 1 (MPM unpubl. data). Although the genus Panaspis is relatively diverse and widely distributed in southern Africa (Medina et al. 2016), the distribution of P cabindae ranges from southwestern Republic of the Congo and the Democratic Republic of the Congo southwards to central and southwestern Angola through woodlands associated with the Angolan Escarp- ment (Ceriaco et al. 2020b). The putative new species Panaspis sp. 1 belongs to
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 177
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Table 3. Comparison of herpetofaunal endemicity levels between Angolan and Namibian inselbergs. Data from Bauer et
al. (2023) and Becker et al. (2023).
Country Inselberg / Highlands
Namibia Brandberg Erongo Mts Spitzkoppe
Swakop-Kahn inselbergs
Central Highlands Tiras Mountains Huns-Orange Mts Baynes-Otjihipa Entedeka Mts Karasberg Klein Karasberg Nubib Mts Aus Mts Otavi Highlands Huab Outliers Brukkaros Onder-Rooirand Interior Plateau Namuskluft Mts south Otjihipa Mts Waterberg Plateau Skerpioenkop
Central Group south to Karasberg exclusive of desert inselbergs
Naukluft Mts Tsaris Mts Gamsberg Rantberge Central Plateau Rooikoppe Auas Mountains Angola Namba Serra da Neve Huila Escarpment and Plateau Central Plateau Pungo Andongo Mt. Moco Mombolo
Congulu Escarpments
Strictly endemic species (Amphibians / Reptiles)
Strictly highland but not strictly endemic (Amphibians / Reptiles)
ihe -/2 ef -/1 a9 = -/1 =f -/3 ze a ae | -/1 -/3 =e -/3 -/1 -/- = ave =f -/3 =f -/1 -/1 -/2 - =f -/2 -/- =/- -/1 = Lite f= -/1 -/1 -/- -/- 1/1 eae Bi -/2 -/- == -/1 ar -/1 =f -/1 wks -/1 ip -/1 a ae -/1 Se -/1 ia -/1 =f -/1 1/1 i= tT 1/4 2/5 1/4 2/1 1/6 -/1 -/- -/1 -/1 ae a
3/- 2
the same Central African lineages as P cabindae, the same clade as the species from the Gulf of Guinea Oceanic islands (Ceriaco et al. 2020b; MPM unpubl. data).
On the other hand, Lygodactylus baptistai belongs to a lineage comprising East African species (Marques et al. 2020). As part of the A. occidentalis species
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750
178
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
complex, a group whose distribution covers most of southern Africa, A. muk- wando is more closely related to A. percivali, a species endemic to the Eastern Arc Mountains of northeastern Tanzania and southeast Kenya, than to other lin- eages of the complex occurring in Namibia and Angola (Marques et al. 2023b). This pattern is also present in the new species Panaspis sp. 2, currently under description (MPM unpubl. data). Among the taxa with southern African affinities there are particularly arid-adapted lizards, which are mostly restricted to the dry lowlands of the inselberg, such as Pedioplanis haackei Conradie, Measey, Branch & Tolley, 2012, Pedioplanis serodioi Parrinha, Marques, Heinicke, Khalid, Parker, Tolley, Childers, Conradie, Bauer & Cerfaco, 2021, Heliobolus crawfordi Marques, Ceriaco, Heinicke, Chehouri, Conradie, Tolley & Bauer, 2022, and Pachydactylus caraculicus Fitzsimons, 1959. On the other hand, the more mesic highlands of the inselberg also support taxa that are typically associated with the highlands of the Escarpment and the Central Plateau, such as Trachylepis ansorgii, Trachylepis huilensis, Psammophylax tritaeniatus (Giinther, 1868), and Leptopelis anchietae.
Taxonomic accounts
Amphibia, Anura Family Pipidae Gray, 1825 Genus Xenopus Wagler, 1827
Xenopus petersii Bocage, 1895 Fig. 4a, b
Records. Catchi, small stream near basecamp [-13.7630, 13.2513, 1595 ml (MUNHAC/MB04-001066); Maylowe, inside a well [-13.8349, 13.2765, 803 ml] (MUNHAC/MB04-001067-001091).
Comments. Traditionally considered as a subspecies of Xenopus laevis (Daudin, 1802), Furman et al. (2015) reclassified all central and western Ango- lan populations as X. petersii. Frétey et al. (2018) designated as lectotype of the species a specimen collected by José d’'Anchieta in Benguela Province, Angola. This species is known from several localities in southwestern regions of the country (see Marques et al. 2018) and the Serra da Neve population is within the expected distribution range of X. petersii in the province.
Family Bufonidae Gray, 1825
Genus Poyntonophrynus Frost, Grant, Faivovich, Bain, Haas, Haddad, de Sa, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler, 2006
Poyntonophrynus grandisonae (Poynton & Haacke, 1993) Fig. 4c
Records. N’Dolondolo [-13.8133, 13.1362, 889 m] (CAS 262731, 262732; UF 184185-184187; INBAC/AMB 10337, 10339).
Comments. These are the first specimens of P. grandisonae collected since the original description by Poynton and Haacke (1993) and used by Ceriaco et al. (2018) to provide the first molecular data for P grandisonae. The natural
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 179
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Figure 4. a, b Xenopus petersii ¢ Poyntonophrynus dombemsis d Poyntonophrynus pachnodes e Sclerophrys pusilla
f Phrynomantis annectens g Leptopelis anchietae h Phrynobatrachus natalensis. Photographs by AT (a, b, e, g, h); LMPC (c, f) and IA (d).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 180
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
history and distribution of this species remain poorly known. The species is endemic to Angola and restricted to the southwestern regions of Namibe Prov- ince. It is associated with xeric vegetation in low-elevation areas (Marques et al. 2018; Baptista et al. 2023).
Poyntonophrynus pachnodes Ceriaco, Marques, Bandeira, Agarwal, Stanley, Bauer, Heinicke & Blackburn, 2018 Fig. 4d
Records. Basecamp 1 [-13.7770, 13.259, 1488 m] (UF 184183, 184184, 190278, 190279; CAS 262730, INBAC/AMB 10209; INBAC/LMPC 1262-1264; MUHNAC/MB04-000999, 001092).
Comments. This recently described species is only known from Serra da Neve and considered a strict endemic (Ceriaco et al. 2018; Baptista et al. 2023). Additional material for this species was collected from the type locality by the second expedition to the inselberg in 2019. This highland specialist was found in an area with moist soil under leaf-litter and rocks, at elevations greater than 1000 m. Recent research by Baptista et al. (2023) provided new data regarding the phylogenetic relationship of this Serra da Neve endemic, indicating that it is closely related to the arid-adapted P dombensis and P. damaranus.
Genus Sclerophrys Tschudi, 1838
Sclerophrys pusilla (Mertens, 1937) Fig. de
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 263036, 263037; IN- BAC/AMB 10206); vic. N’Dolondolo [-13.8087, 13.1352, 731 m] (CAS 263025, 263030); N’Dolondolo [-13.8133, 13.1362, 593 m] (UF 187174); Maylowe [-13.8342, 13.2767, 803 m] (UF 190289; MUNHAC/MB04-001093); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MB04-001094).
Comments. The species has a wide distributional range in central and south- ern Angola (Marques et al. 2018). The Angolan population was for some time treated as Sclerophrys maculata (Hallowell, 1854), until Poynton et al. (2016) established that S. maculata is restricted to West Africa, while S. pusilla is main- ly distributed across eastern and southern Africa, including Angola. This spe- cies is common in Serra da Neve and its surroundings. It is usually found near leaf litter.
Family Microhylidae Gunther, 1858 Genus Phrynomantis Peters, 1867
Phrynomantis annectens Werner, 1910 Fig. 4f
Records. N’Dolondolo [-13.8004, 13.1362, 897 m] (UF 187250, 187251; INBAC/ AMB 10344); Maylowe [-13.8342, 13.2767, 803 m] (UF 190275).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 181
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Comments. Endemic to southwestern Africa from southwestern Ango- la southwards through Namibia to the arid regions of northern South Africa (Marques et al. 2018; Channing and Rédel 2019). In Angola, P annectens is restricted to the coastal lowlands of the country (Marques et al. 2018). The species is frequently associated with inselbergs and other rocky outcrops in arid regions (Channing 2001; Channing and Rédel 2019). Our specimens were found inside crevices at the base of Serra da Neve. Ceriaco et al. (2021) dis- cussed the taxonomic and nomenclatural history of P annectens and provided an additional record for the country.
Family Arthroleptidae Mivart, 1869 Genus Leptopelis Gunther, 1859
Leptopelis anchietae (Bocage, 1873) Fig. 4g
Records. Catchi, near small stream [-13.7630, 13.2514, 1595 m] (MUNHAC/ MB04- 001095, 001096).
Comments. An Angolan endemic, distributed throughout much of the west- ern half of the country (Marques et al. 2018; Becker et al. 2023). The collected specimens were heard croaking from a small bush at night, a common event especially after rain. Considered a cryptic species complex by several authors (Perret 1976; Schigtz 1999). Becker et al. (2023) did not record the species in Serra da Neve.
Family Ptychadenidae Dubois, 1987 Genus Ptychadena Boulenger, 1917
Ptychadena anchietae (Bocage, 1867)
Records. N’Dolondolo [-13.8004, 13.1362, 897 m] (CAS 263024; UF 187280; INBAC/AMB 10308).
Comments. The species is broadly distributed from western to eastern An- gola and is widespread extralimitally (Marques et al. 2018; Channing and Rodel 2019), typically associated with savanna and grassland habitats.
Family Phrynobatrachidae Laurent, 1941 Genus Phrynobatrachus Gunther, 1862
Phrynobatrachus natalensis (Smith, 1849) Figs 4e, 5a
Records. Mamué riparian area [-13.8004, 13.1246, 732 m] (UF 187249): Catchi, small stream near basecamp [-13.7630, 13.2513, 1595 m] (MUNHAC/MB04- 001097-001146).
Comments. The species, as currently understood, is likely to contain multiple undescribed cryptic species across its large distribution in the savanna and grassland regions of sub-Saharan Africa (Zimkus et al. 2010). Based on newly
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 182
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
generated molecular data (LMPC unpub. data), we conclude that the population from Serra da Neve belongs to the same lineage as topotypical populations in South Africa. The specimens were collected on the margins of small streams, both in the lowlands and highlands of the inselberg.
Family Pyxicephalidae Bonaparte, 1850 Genus Amietia Dubois, 1987
Amietia angolensis (Bocage, 1866) Fig. 5b
Records. Catchi, small stream near basecamp [-13.7630, 13.2513, 1595 ml (MUNHAC/MB04-001147-001154).
Comments. Formerly considered a widespread species in western and cen- tral Africa, is now restricted to Angola (Channing and Baptista 2013; Channing et al. 2016). The species was frequently encountered in the rocky margins of small streams and ponds at the top of the inselberg.
Genus Tomopterna Dumeril & Bibron, 1841
Tomopterna ahli (Deckert, 1938) Fig. 5c
Record. Maylowe [-13.8342, 13.2767, 803 m] (UF 190305).
Comments. Ceriaco et al. (2016) provided the first record of this species for Angola as Tomopterna damarensis Dawood & Channing, 2002. Heinicke et al. (2017a) used morphological and mtDNA data to show that this species is broadly distributed in Angola and Namibia and suggested that some records of Tomopterna from southern Angola could correspond to this species. Chan- ning and Becker (2019) showed that Tomopterna ahli was a senior synonym of T. damarensis. All Angolan specimens previously assigned to T. damarensis (Ceriaco et al. 2016, Heinicke et al. 2017a; Marques et al. 2018) should thus be referred to as T. ahi.
Tomopterna tuberculosa (Boulenger, 1882) Fig. 5d
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 263038, 263039); Mamweé riparian area [-13.8006, 13.1230, 706 m] (UF 187293); N’Dolondolo [-13.8133, 13.1362, 681 m] (UF 187294); Maylowe [-13.8342, 13.2767, 803 m] (MUNHAC/MB04-001155); Catchi, basecamp [-13.7627, 13.2562, 1597 m] (MUNHAC/MB04-0011556).
Comments. The species has a wide distribution in western Angola (Marques et al. 2018). These specimens represent the first record of the species in Na- mibe Province since those that were reported by Bocage (1895). Tomopter- na tuberculosa appears to occur in sympatry with T. ahli at Serra da Neve and across the southwestern areas of Namibe Province.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 183
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Figure 5. a Phrynobatrachus natalensis b Amietia angolensis c Tomopterna ahli d Tomopterna tuberculosa e Kinixys belli- ana f Stigmochelys pardalis g Afroedura praedicta h Hemidactylus benguellensis. Photographs by AT (a, b, d, h) and LMPC
(c, e-g).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 184
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Reptilia, Testudines Family Testudinidae Batsch, 1788 Genus Kinixys Bell, 1827
Kinixys belliana Gray, 1863 Fig. 5e
Records. Catchi surroundings [-13.7577, 13.2543, 1576 m] (MUNHAC/MB03- 001548, 001549 only tissue); 2 km N of Maylowe [-13.8280, 13.2625, 820 ml] (MUNHAC/MB03-001550, only tissue).
Comments. A wide-ranging species from eastern Africa to southwestern and central Angola. Historically, the taxonomic status of Kinixys populations in Angola has been uncertain (for further discussion see Marques et al. 2018), although according to the most recent sub-Saharan Africa chelonian phylogeny (Fritz et al. 2022), the Angolan material should be assigned to Kinixys belliana. This species is frequently consumed a delicacy by the Mucubal tribe at the base of Serra da Neve (pers. obs.).
Genus Stigmochelys Gray, 1873
Stigmochelys pardalis (Bell, 1828) Fig. 5f
Record. 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MBO3- 001551, only tissue).
Comments. Stigmochelys pardalis is a large-bodied species occurring from northern Somalia southwards through eastern Africa to South Africa, and westwards to Namibia and Angola (Marques et al. 2018). The species has been recorded in the southern provinces of the country (Marques et al. 2018).
Reptilia, Squamata Family Gekkonidae Gray, 1825 Genus Afroedura Loveridge, 1944
Afroedura praedicta Branch, Schmitz, Lobon-Rovira, Baptista, Antonio & Conradie, 2021 Fig. 5g
Records. Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 ml (MUNHAC/ MB03-001552-001554).
Comments. This recently described species is only known from the Serra da Neve inselberg and is considered a strict endemic (Branch et al. 2021; Conradie et al. 2023). This highland specialist was found in vertical walls of large granite boulders at night. A juvenile (MUNHAC/ MB03-001552) was collected in rock crevices near a communal laying site with dozens of hatched eggs.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 185
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Genus Hemidactylus Goldfuss, 1820
Hemidactylus benguellensis Bocage, 1893 Fig. 5h
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 263367—263372); N’Do- londolo [-13.8133, 13.1362, 681 m] (CAS 263536-263540; UF 187202; INBAC/ AMB 10237, 10245); vic. N’Dolondolo [-13.8068, 13.1351, 754 m] (CAS 263549); Maylowe [-13.8342, 13.2767, 803 m] (CAS 266144); Catchi, rock outcrops near ba- secamp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001555, 001556); MPLA post near Catchi [-13.7618, 13.2514, 1614m] (MUNHAC/MB03-001799); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MB03-001557, 001558).
Comments. The species is endemic to southwestern Angola and northern Namibia (Marques et al. 2018; Ceriaco et al. 2020c; Lob6n-Rovira et al. 2021). Although H. benguellensis appears to be very common in the Angolan Escarp- ment areas, it has also been found in more coastal environments in southwest- ern Namibe Province (Lob6n-Rovira et al. 2021). The species is strongly asso- ciated with rupiculous habitats.
Genus Chondrodactylus Peters, 1870
Chondrodactylus pulitzerae (Schmidt, 1933) Fig. 6a
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 266367-CAS 266369, 266371; INBAC/AMB 10200); N’Dolondolo [-13.8133, 13.1362, 681 m] (CAS 266370); Maylowe [-13.8342, 13.2767, 803 m a.s.l] (CAS 266114); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03- 001559- 001565); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/ MB03-001566-001 568).
Comments. The genus has recently been reviewed by Heinz et al. (2021). The species is quite widespread in Angola (Marques et al. 2018; Heinz et al. 2021), and it occurs at different elevations on Serra da Neve and its surrounding areas.
Genus Lygodactylus Gray, 1864
Lygodactylus baptistai Marques, Ceriaco, Buehler, Bandeira, Janota & Bauer, 2020 Fig. 6b
Records. Mamué riparian area [-13.8008, 13.1235, 715 m] (CAS 263557), [-13.8004, 13.1246, 748 m] (CAS 263551); MPLA post near Catchi [-13.7618, 13.2514, 1614 m] (MUNHAC/MB03-001569-001571); near Ondjili, between Catchi and Lutala crater (MUNHAC/MB03-001572).
Comments. This recently described species is only known from the Serra da Neve inselberg and is considered a strict endemic. The species was recently described from the inselberg by Marques et al. (2020). Lygodactylus baptistai appears to be the single representative of this lineage in southwestern Africa. Morphologically it is also more similar to those species found on inselbergs of
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 186
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
(adult and juvenile) f Pachydactylus maiatoi g Pachydactylus cf. punctatus h Rhoptropus aff. barnardi. Photographs by AT (a, d, e, g); IA (b, f) and LMPC (ce, h).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 187
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Mozambique such as L. rex Broadley, 1963 and L. regulus Portik, Travers, Bau- er & Branch, 2013 (Portik et al. 2013b) than to the other species known from Angola, L. angolensis Bocage, 1896 and L. nyaneka Marques, Ceriaco, Buehler, Bandeira, Janota & Bauer, 2020, both restricted to Miombo forested areas, or L. lawrencei Hewitt, 1926, an arid zone specialist.
Lygodactylus nyaneka Marques, Ceriaco, Buehler, Bandeira, Janota & Bauer, 2020 Fig. 6c
Record. Mamué riparian area [-13.8008, 13.1235, 715 m] (CAS 263556).
Comments. A recently described species from the central and southwestern regions of the country and neighboring northern Namibia, with records from Epupa Falls (Marques et al. 2020). The individual collected from Serra da Neve occurs in sympatry with the strictly endemic L. baptistai.
Genus Pachydactylus Wiegmann, 1834
Pachydactylus caraculicus FitzSimons, 1959 Fig. 6d,e
Records. N’Dolondolo [-13.8133, 13.1362, 681 m] (CAS 10283, 10347); May- lowe [-13.8342, 13.2767, 803 m] (CAS 266145; MUNHAC/MB03-001573); 2 km N of Maylowe [-13.8280, 13.2616, 804 m] (MUNHAC/MB03-001574); 2 km N of Maylowe, rock outcrops near basecamp [-13.8280, 13.2646, 8020 m] (MUN- HAC/MB03-001575-001581).
Comments. The species is known from southwestern Angola and northwest- ern Namibia (Marques et al. 2018). It is part of a diverse and primarily rupiculous “northwestern clade” of Pachydactylus (Bauer and Lamb 2005; Heinicke et al. 2011), sister to P angolensis Loveridge, 1944 and P. maiatoi (Heinicke et al. 2017b; Marques et al. 2023a). Usually found in crevices and cracks of granitic boulders.
Pachydactylus maiatoi Marques, Parrinha, Ceriaco, Brennan, Heinicke & Bauer, 2023 Fig. 6f
Records. N’Dolondolo [-13.8133, 13.1362, 681 m] (CAS 266484, 266485); Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 266486); Maylowe [-13.8355, 13.2755, 798 m] (MUNHAC/MB03-001246); Catchi, rock outcrops near base- camp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001247); 2 km N of May- lowe [-13.8289, 13.2625, 820 m] (MUNHAC/MB03-001248).
Comments. Heinicke et al. (2017b) identified putative species-level diversity within the Pachydactylus angolensis group. Based on a combination of mor- phological, meristic and DNA sequence data, it was possible to separate two different taxa, a “coastal” and “inland” form of P angolensis group. Marques et al. (2023a) described the inland form as P maiatoi. The recently described species appears to be restricted to southwestern Angola, namely in the inland
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 188
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
regions of Namibe Province and along the highlands associated with the Es- carpment in Huila Province. This species is usually found under rocks in areas with some vegetation, in highland regions.
Pachydactylus cf. punctatus Peters, 1854 Fig. 6g
Records. 2 km N of Maylowe [-13.8289, 13.2625, 820 m] (MUNHAC/MB03- 001582- 001591).
Comments. Pachydactylus punctatus is a widespread species complex ex- tending from South Africa northwards to Malawi, the former Katanga Province of the Democratic Republic of the Congo and southern Angola (Marques et al. 2018). In Angola itself, Heinz (2011) noted the presence of four species-level divergent lineages. As the taxonomy and nomenclature of this group is still ina state of flux, we opt here to simply refer our specimens to P. cf. punctatus, until the ongoing revision of the group is complete.
Genus Rhoptropus Peters, 1869
Rhoptropus aff. barnardi Hewitt, 1926 Fig. 6h
Records. N’Dolondolo [-13.8133, 13.1316, 681 m] (MUNHAC/MB03-001592- 001595); vic. N’Dolondolo [-13.8113, 13.1365, 699 m] (MUNHAC/ MBO03- 001596-001603); Basecamp 1 [-13.7770, 13.2591, 1488 m] (MUNHAC/MB03- 001604-001610); Maylowe [-13.8355, 13.2755, 798 m] (CAS 266105-10, 266130-32, 266134-37, 266156; MUNHAC/MBO03-001 611-001 620); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 m] (CAS 266127; MUNHAC/ MB03-001621, 001622); near Basecamp 1 [-13.7770, 13.2591, 1488 m] (MUN- HAC/ MB03-001623-001633; CAS 266164—68); Catchi surroundings [-13.7619, 13.2568, 1585 m] (MUNHAC/MB03-001634-001688); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001689-001 693); MPLA post near Catchi [-13.7618, 13.2514, 1614 m] (MUNHAC/MB03-001 694); 2 km N of Maylowe [-13.8289, 13.2625, 820 m] (MB03-001695-001705). Comments. Recent surveys in southwestern Angola revealed undescribed cryptic diversity associated with Rhoptropus barnardi (Ceriaco et al. 2016; Kuhn 2016; Butler et al. 2019; Lob6n-Rovira et al. 2022). The specimens collected in Serra da Neve belong to an undescribed Angolan endemic with affinities to Rhoptropus barnardi Hewitt, 1926 and R. biporosus FitzSimons, 1957, which is widespread in southwestern Angola, from sea level to an elevation of more than 2000 m. A revision of the genus is being prepared (DP unpubl. data).
Rhoptropus aff. montanus Laurent, 1964 Fig. /a
Records. Catchi surroundings [-13.7618, 13.2514, 1614 m] (MUNHAC/MB03- 001706-001708).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 189
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Comments. Three specimens of an unknown species of Rhoptropus were collected in a riparian area during our last expedition in 2022. This lineage is currently being described in a separate paper, and is endemic to the highlands of the inselberg (DP unpubl. data). It has affinities to Rhoptropus montanus Lau- rent, 1964, another montane endemic from the Huila Plateau in southwestern Angola (Laurent 1964).
Family Lacertidae Oppel, 1811 Genus Heliobolus Fitzinger, 1843
Heliobolus crawfordi Marques, Ceriaco, Heinicke, Chehouri, Conradie, Tolley & Bauer, 2022 Fig. 7b, c
Records. N’Dolondolo [-13.8133, 13.1316, 681 m] (CAS 266267-266269, 266271, 266273, 266275; INBAC/AMB 10335); near Maylowe [-13.8113, 13.3222, 879 m] (CAS 266120, 266121, 266146; INBAC/LMPC 1220, 1221; MHNC-UP/REP 869-871); 2 km N of Maylowe [-13.8280, 13.2416, 804 ml (MUNHCA/MB03-001709-001713).
Comments. Endemic to Angola and restricted to the central coastal regions of the country, including the surroundings of Serra da Neve (Marques et al. 2022). It is absent from higher elevations in Serra da Neve but abundant in the surrounding lowlands.
Genus Pedioplanis Fitzinger, 1843
Pedioplanis haackei Conradie, Measey, Branch & Tolley, 2012 Fig. 7d
Records. Dirt road to the top of the mountain, near Maylowe [-13.8328, 13.2652, 794 m] (MHNC-UP/REP 634-635, CAS 266123); Dirt road to Quilengues [-13.8159, 13.3264, 892 m] (MHNC-UP/REP 645); vic. N’Dolondolo [-13.8086, 13.13521, 731 m] (CAS 264757, 264765); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MB03-001714-001716).
Comments. This species is endemic mainly to Namibe Province, usually oc- curring near granite outcrops on sandy areas (Conradie et al. 2012; Parrinha et al. 2021). It was only found in the lowland arid plains of Serra da Neve surroundings, in Mopane habitats, but not in the Miombo-dominated areas at higher altitudes.
Pedioplanis serodioi Parrinha, Marques, Heinicke, Khalid, Parker, Tolley, Childers, Conradie, Bauer & Ceriaco, 2021 Fig. /e
Records. Dirt road to the top of the mountain, near Maylowe [-13.8328, 13.2652, 794 m] (MHNC-UP/REP 636; CAS 266122); Dirt road to Quilengues [-13.8159, 13.3264, 892 m] (MHNC-UP/REP 637, 638); Maylowe [-13.8355, 13.2755, 798 m] (MHNC-UP/REP 639, 646); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MB03-001717-001756).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 190
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Figure 7. a Rhoptropus aff. montanus b Heliobolus crawfordi (adult) ec Heliobolus crawfordi (juvenile) d Pedioplanis haackei e Pedioplanis serodioi f Cordylus phonolithos g Cordylosaurus subtessellatus h Gerrhosaurus sp. Photographs by AT (a, b, e, f, h) and LMPC (ce, d, g).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 191
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Comments. This recently described species (Parrinha et al. 2021) is widely distributed through the lowlands of southwestern Angola, from central Ben- guela Province to western Cunene Province, with exception of the more xeric areas of southwestern Namibe Province. As with other lacertids, this species was only recorded from the arid lowlands of the inselberg.
Family Cordylidae Fitzinger, 1826 Genus Cordylus Laurenti, 1768
Cordylus phonolithos Marques, Ceriaco, Stanley, Bandeira, Agarwal & Bauer, 2019 Fig. 7f
Records. vic. N’Dolondolo [-13.8068, 13.1351, 752 m] (CAS 263581; INBAC/ AMB 10272); Rock outcrops near Catchi [-13.7653, 13.2571, 1645 m] (MUN- HAC/MB03-001757-001765).
Comments. This recently described species is only known from the Serra da Neve inselberg and is considered a strict endemic. Cordylus phonolithos was recently described from the inselberg by Marques et al. (2019). It is genetically divergent and morphologically distinguished from the closely related Angolan Escarpment dwelling Cordylus machadoi Laurent, 1964 and the low-elevation species C. namakuiyus Stanley, Ceriaco, Bandeira, Valério, Bates & Branch, 2016. This species is found in cracks in granite boulders, but sometimes can be seen basking outside or even crossing paths on the ground.
Family Gerrhosauridae Fitzinger, 1843 Genus Cordylosaurus Gray, 1866
Cordylosaurus subtessellatus (Smith, 1844) Fig. 7g
Records. vic. Dolondolo [-13.8087, 13.1352, 731 m] (CAS 263031); Mamué ri- parian area [-13.8003, 13.1229, 710 m] (INBAC/AMB 10326).
Comments. The species is known from southwestern Angola through west- ern Namibia and into western parts of South Africa (Marques et al. 2018). In Angola, C. subtesselatus has been recorded from the coastal areas of Benguela and Namibe provinces (Marques et al. 2018). It is commonly found basking in granite outcrops and hidden in crevices.
Genus Gerrhosaurus Wiegmann, 1828
Gerrhosaurus sp. Fig. 7h
Records. Catchi surroundings [-13.7577, 13.2543, 1576 m] (MUNHAC/MB03- 001766-001778).
Comments. The Serra da Neve population belongs to a genetic clade that also occurs in the plateau areas of central and southeastern Angola, already signaled by Butler et al. (2019) and Conradie et al. (2016b, 2022) (LMPC unpub. data).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 192
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Genus Matobosaurus Bates & Tolley, 2013
Matobosaurus maltzahni (De Grys, 1938) Fig. 8a
Records. N’Dolondolo [-13.8133, 13.1362, 681 m] (INBAC/AMB 10280); Rock outcrops near Catchi [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001779); 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MUNHAC/MB03-001780).
Comments. The species is known from the northwestern regions of Namibia to the southwestern regions of Angola, where it occurs in the western lowlands of Namibe and Benguela provinces and neighboring Huila Province (Marques et al. 2018). It occurs in a variety of micro-habitats, from rock outcrops to woodlands. The specimen MUNHAC/MB03-001780 was found at night, sleep- ing inside a hollow trunk of a Mopane tree.
Family Scincidae Genus Acontias Cuvier, 1816 “1817”
Acontias mukwando Marques, Parrinha, Tiutenko, Lopes-Lima, Bauer & Ceriaco, 2023 Fig. 8b
Records. Catchi, Miombo woodland near basecamp [-13.7660, 13.2587, 1674 m] (MUHNAC/MB03-001522-24).
Comments. This recently described species is only known from the Serra da Neve inselberg and considered a strict endemic. It was found hiding under rocks and active on leaf-litter in Miombo-dominated landscapes (Marques et al. 2023b).
Genus Mochlus Gunther, 1864
Mochlus sundevallii (Smith, 1849)
Record. 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (specimen not collected).
Comments. This species occurs throughout southern Africa (Marques et al. 2018). The specimen was seen active at dusk, under a cattle fence made from dead Mopane trees, in sandy soil with some Mopane leaf litter.
Genus Panaspis Cope, 1868
Panaspis cabindae (Bocage, 1866)
Records. Mamué riparian area [-13. 8015, 13.1206, 665 m] (CAS 263550, 263553-263555; UF 187242; INBAC/AMB 10317, 10320).
Comments. The distribution area of this species extends from central Africa to the central highlands in Angola, reaching its southern limit in the forest margins below the Escarpment in southeastern Namibe Province (Ceriaco et al. 2020b). The species appears to be absent from more xeric and desertic areas of the southwestern regions of Namibe Province, where it is replaced by its congener Panaspis mocamedensis. |In Serra da Neve, the species was found under leaf-lit-
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 193
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Figure 8. a Matobosaurus maltzahni b Acontias mukwando c Panaspis sp. 1 d Sepsina copei e Trachylepis albopunctata f Trachylepis ansorgii g Trachylepis binotata h Trachylepis bouri. Photographs by AT (a-c, e-h); LMPC (d).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 194
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
ter, in dense Miombo woodlands. In Serra da Neve the species occurs in more hu- mid lowlands, tendentially preferring more forested areas with deeper leaf litter.
Panaspis mocamedensis Ceriaco, Heinicke, Parker, Marques & Bauer, 2020
Records. 2 km N of Maylowe [-13.8280, 13.2625, 820 m] (MB03-001532, 001533).
Comments. P mocamedensis is endemic to Namibe Province (Ceriaco et al. 2020b). This species tendentially prefer more open and dry micro-habitats than their Angolan congeners.
Panaspis sp. 1 Fig. 8c
Records. Rock outcrops near Catchi [-13.7653, 13.2571, 1645 m] (MUNHAC/ MB03-001525, 001526); Catchi, basecamp [-13.7627, 13.2562, 1597 m] (MUN- HAC/MB03-001528); 2 km N of Maylowe [-13.8280, 13.2625, 820 ml], (MUN- HAC/MB03-001529-001531); Dry riverbed, 2 km N of Maylowe [-13.8265, 13.2601, 720 m] (MUNHAC/MB03-001534).
Comments. A new species is currently being described (MPM unpubl. data), only known from the Serra da Neve inselberg, where it is assumed to be endem- ic. Panaspis sp. 1 belongs to the same Central African lineages as P cabindae and is part of the same clade as the Gulf of Guinea Oceanic islands species (Ceriaco et al. 2020b; MPM unpubl. data). This species shows some ecological adaptability occurring in both Miombo woodlands in the higher elevation areas but also in the arid Mopane lowlands (MPM unpubl. data). It was recorded in sympatry with P mocamedensis in the lowlands of the inselberg.
Panaspis sp. 2
Record. MPLA post near Catchi [-13.7618, 13.2514, 1614 m] (MUNHAC/MB03- S27}
Comments. A new species is currently being described (MPM unpubl. data); similarly to Panaspis sp. 1, it is known only from Serra da Neve. The single col- lected specimen was found under a long near to a riparian gallery. Panaspis sp. 2 present phylogenetic and biogeographic affinities with P annettesabinae from the highlands of Ethiopia (Colston et al. 2020; MPM unpubl. data).
Genus Sepsina Bocage, 1866
Sepsina copei Bocage, 1873 Fig. 8d
Records. Mamué riparian area [-13.8015, 13.1206, 665 m] (CAS 263918-263921); vic. N'Dolondolo [-13.8086, 13.1352, 731 m] (CAS 263916).
Comments. This species is endemic to Angola. It occurs in western coastal regions, from Luanda to Namibe Province.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 195
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Genus Trachylepis Fitzinger, 1843
Trachylepis albopunctata (Bocage, 1867) Fig. 8e
Records. Rock outcrop near Basecamp 1 [-13.7864, 13.2575, 1596 m] (CAS 263560); Basecamp 1 [-13.7770, 13.2591, 1488 m] (MUNHAC/MB03-001384, 001516; INBAC/LMPC 1265); Catchi surroundings [-13.7620, 13.2569, 1585 ml] (MUNHAC/MB03-001468, 001469, 001486-001498, 001500-001502, 001504-— 001508); 2 km N of Maylowe [-13.8280, 13.2625, 818 m] (MUHNAC/MBO03- 001509); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 ml (MUNHAC/MB03-001499, 001503).
Comments. The species is widely distributed in Angola, except for the south- eastern areas of the country, where it is replaced by Trachylepis damarana (Pe- ters, 1870). The Angolan population was long identified as Trachylepis varia (Peters, 1867), but recent reviews by Weinell and Bauer (2018) and Ceriaco et al. (2024) validated the specific status of albopunctata.
Trachylepis ansorgii (Boulenger, 1907) Fig. 8f
Records. Maylowe [-13.8357, 13.2763, 800 m] (MUHNAC/MBO03-001396, 001397); Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 263567; UF 187313); Rock outcrop near Basecamp 1 [-13.7865, 13.2572, 1594 m] (UF 187314); vic. N’Dolondolo [-13.8104, 13.1361, 713 m] (CAS 263545); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001472, 001475, 001477, 001478, 001482); Catchi surroundings [-13.7620, 13.2569, 1585 ml (MUNHAC/MB03-001473, 001474, 001479-001481, 001485); 2 km N of May- lowe [-13.8280, 13.2625, 818 m] (MUHNAC/MB03-001476); Lutala crater, near airstrip [-13.7325, 13.1841, 1567 m] (MUHNAC/MB03-1484).
Comments. Butler et al. (2019) presented the first topotypical material since the taxon was originally described by Boulenger (1907). According to the mo- lecular results of Butler et al. (in press) and Ceriaco et al. (2024), T. ansorgii is a valid species, distinct from T. sulcata. Trachylepis ansorgii is restricted to the highlands of Angola, mostly distributed from Malanje to northern Huila Prov- ince, while T. sulcata seems to occur in the southwestern regions of the coun- try, with records for southern Namibe and southwestern Huila provinces (Ceria- co et al. 2024). The material collected from Serra da Neve is the first evidence of T. ansorgii in Namibe Province and the southernmost record of the species, emphasizing the species restriction to areas of high elevations.
Trachylepis binotata (Bocage, 1867)
Fig. 8g
Records. Maylowe [-13.8357, 13.2763, 800 m] (CAS 266149); 2 km N of May- lowe [-13.8280, 13.2625, 818 m] (MUHNAC/MB03-001453, 001454).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 196
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Comments. This large arboreal skink occurs across all southwestern Ango- la. Trachylepis binotata is usually associated with Mopane woodland habitats and was widely recorded in the country (Marques et al. 2018; Ceriaco et al. 2024). In our study area, it was only found in the Mopane-dominated localities at the base of the inselberg.
Trachylepis bouri Ceriaco, Marques, Parrinha, Tiutenko, Weinell, Butler & Bauer, 2024 Fig. 8h
Records. Catchi surroundings [-13.7620, 13.2569, 1585 m] (MUNHAC/MB03- 001511); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 ml (MUNHAC/MB03-001 512).
Comments. This newly described species is endemic to southwestern Ango- la. It is associated with rock outcrops, especially along the Escarpment (Ceria- co et al. 2024).
Trachylepis chimbana (Boulenger, 1887) Fig. 9a
Records. vic. N’Dolondolo [-13.8105, 13.1361, 707 m] (CAS 263542, 263543, 263544); near dirt road to top of the mountain, N of Maylowe [-13.8105, 13.2581, 1502 m] (CAS 263562, 263563); Maylowe [-13.8355, 13.2755, 798 ml] (MUNHAC/MB03-001387); 2 km N of Maylowe [-13.8280, 13.2625, 818 ml (MUHNAC/MB03-001518); Catchi surroundings [-13.7619, 13.2569, 1585 ml] (MUNHAC/MB03-001514); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 m] (MUNHAC/MB03-001515, 001519).
Comments. Trachylepis chimbana is one of the most taxonomically chal- lenging species of the genus. This Angolan endemic had been confused by sev- eral authors (Schmidt 1933; Hellmich 1957a, b; Laurent 1964) with other taxa, such as Trachylepis bocagii or T. wahlbergi. Ceriaco et al. (2024) restricted T. chimbana to northern Namibe and southern Benguela provinces. This species is primarily rupicolous and was found mainly on granite outcrops.
Trachylepis huilensis (Laurent, 1964) Fig. 9b
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (CAS 263565; UF 187310; MUNHAC/MB03-001388, 001516); Rock outcrop near Basecamp 1 [-13.7877, 13.2572, 1600 m] (UF 187311), [-13.7865, 13.2572, 1594 m] (CAS 263558), [-13.7881, 13.2571, 1612 m] (CAS 263559); Catchi surroundings [-13.7619, 13.2569, 1585 m] (MUNHAC/MB03-001467, 001470, 001471, 001517). Comments. This taxon was originally described as a subspecies of T. bayonii by Laurent (1964). Butler et al. (2019) provided the first evidence that huilensis should be considered a full species, rather than a subspecies. Based on newly
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 197
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
obtained molecular and morphological data, Ceriaco et al. (2024) found that the species is more widespread than originally known. It is associated with the highlands of the Leba Escarpment in Huila Province and the Serra da Neve inselberg.
Trachylepis laevis (Boulenger, 1907) Fig. 9c
Records. vic. N’Dolondolo [-13.8105, 13.1361, 713 m] (CAS 263541; UF 187308); Rock outcrop near Basecamp 1 [-13.7881, 13.2571, 1612 m] (CAS 263582); vic. Catci, granite boulders near basecamp [-13.7646, 13.2601, 1603 m] (MUHNAC/ MB03-001463-001465).
Comments. This conspicuous species is common in southwestern Angola and neighboring Namibia (Marques et al. 2018; Ceriaco et al. 2024). It is usually seen basking on granite outcrops.
Family Chamaeleonidae Rafinesque, 1815 Genus Chamaeleo Linnaeus, 1758
Chamaeleo dilepis Leach, 1819 Fig. 9d
Records. Catchi surroundings [-13.7620, 13.2569, 1585 m] (MUNHAC/MB03- 001781-001782).
Comments. Chamaeleo dilepis is a species complex distributed throughout southern and eastern Africa (Tilbury 2010). Marques et al. (2018) opted to iden- tify all Angolan records as C. dilepis quilensis and showed its large distribution throughout the country. More recently, Main et al. (2022) provided evidence of several divergent lineages that may warrant a species-level status. For this pa- per and until the taxonomy of the group is clarified, we simply refer to the Serra da Neve populations as C. dilepis.
Family Agamidae Gray, 1827 Genus Agama Daudin, 1802
Agama aculeata Merrem, 1820 Fig. 9e
Records. Maylowe [-13.8342, 13.2767, 803 m] (CAS 266138, 266139).
Comments. Agama aculeata is a ground-dwelling agamid commonly found in higher altitudes in southern Angola (Marques et al. 2018). In Namibe Prov- ince the species is restricted to areas closely associated with inselbergs and with the Great Escarpment, being replaced by A. anchietae in lower altitudes (Marques et al. 2018).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 198
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Figure 9. a Trachylepis chimbana b Trachylepis huillensis ¢ Trachylepis laevis d Chamaeleo dilepis e Agama aculeata f Agama schacki (male) g Agama schacki (female) h Afrotyphlops schlegeli petersii. Photographs by AT (a, c, d, f, g) and LMPC (b, e, h).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 199
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Agama schacki Mertens, 1938 Fig. 9f, g
Records. Rock outcrop near Basecamp 1 [-13.7865, 13.2572, 1594 ml] (CAS 263035), [-13.7881, 13.2571, 1612 m] (INBAC/AMB 10233); N’Dolondo- lo [-13.8133, 13.1362, 681 m] (INBAC/AMB 10302, 10303); vic. N’Dolondo- lo [-18.8105, 13.1361, 707 m] (CAS 263026); Maylowe [-13.83424, 13.27669, 803 m] (CAS 266128, CAS 266129; INBAC/LMPC 1169; MUNHAC/MB03- 001783); Catchi surroundings [-13.7620, 13.2569, 1584 m] (MUNHAC/MB03- 001784); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 ml (MUNHAC/MB03-001785-001796); 2 km N of Maylowe [-13.8280, 13.2616, 804 m] (MUNHAC/MB03-001797); Serra da Neve base, 2 km N of Maylowe, rock outcrops near basecamp [-13.8280, 13.2625, 820 m] (MUNHAC/MB03-001798).
Comments. Originally described by Mertens (1938) as a subspecies of Ag- ama planiceps Peters, 1862, A. schacki was shown to be a distinct species by Butler et al. (in press), based on molecular data. Marques et al. (2018) had already considered A. schacki as a full species and restricted its distribution to the Escarpment area in western Huila and eastern Namibe and Benguela prov- inces. The species is strongly associated with higher elevations, being replaced by A. planiceps in lowland areas of southern Namibe Province.
Family Typhlopidae Genus Afrotyphlops Broadley & Wallach, 2009
Afrotyphlops schlegeli petersii (Bocage, 1873) Fig. 9h
Record. Mamué riparian area [-13.8015, 13.1206, 665 m] (CAS 266467).
Comments. The Afrotyphlops schlegeli species complex is, as most of the other members of this genus, a taxonomic and nomenclatural conundrum. The validity of petersii as a distinct taxon, endemic to southwestern Angola and Namibia, was supported by Roux-Estéve (1974). Marques et al. (2018) considered the Angolan population simply as A. schlegeli. Given the significant geographic separation between the Angolan population and the topotypical population in Mozambique, as well as the morphological differences noted by Roux- Estéve (1974), we treat the Angolan populations as a distinct subspe- cies until a more comprehensive review of the group is undertaken. This taxon appears to be associated with higher elevation and montane areas (Marques et al. 2018). The collected specimen had been killed by locals who regard it as highly venomous.
Family Leptotyphlopidae Genus Leptotyphlops Fitzinger, 1843
Leptotyphlops cf. scutifrons (Peters, 1854)
Record. vic. Mamué [-13.7877, 13.1257, 1600 m] (CAS 264756). Comments. Leptotyphlops scutifrons is widely distributed in southern Africa. It was recorded from several localities in the Angolan Plateau (Broadley and
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 200
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Broadley 1999; Conradie et al. 2016b; Marques et al. 2018). Although some historical records may refer to Namibiana latifrons (Sternfeld, 1908) (Marques et al. 2018), our specimen agrees almost completely with the diagnosis of L. scutifrons that was provided by Broadley and Broadley (1999). Nevertheless, this taxon should be treated as a species complex with multiple divergent lin- eages (Adalsteinsson et al. 2009). In Angola, it is mostly restricted to the high- lands of the Escarpment and Central Plateau (Marques et al. 2018).
Family Pythonidae Fitzinger, 1826 Genus Python Daudin, 1803
Python natalensis Smith, 1833 Fig. 10a
Record. Maylowe [-13.8342, 13.2767, 803 m] (photographic record, specimens not collected).
Comments. This species is widely distributed throughout southern Angola (Marques et al. 2018). The skin in the photograph is held by local Mucubal peo- ple in the town of Maylowe, where it was killed.
Family Viperidae Oppel, 1811 Genus Bitis Gray, 1842 Subgenus Macrocerastes Reuss, 1939
Bitis (Macrocerastes) gabonica (Dumeéril, Bibron & Duméril, 1854) Fig. 10b
Record. Catchi surroundings [precise locality unknown] (MUNHAC/MBO3- 001535).
Comments. The Gaboon viper is an African viperid with a primarily west and central African distribution, while some populations extend marginally into northern Zambia. Isolated populations exist from southern South Sudan, Ken- ya, Tanzania, and the KwaZulu-Natal region of eastern South Africa (Spawls et al. 2023; Marques et al. 2018). In Angola, the species has been recorded mainly in northern and central areas of the country, with the southwestern-most record in Central Benguela (Marques et al. 2018). The Serra da Neve population thus represents the southwestern-most record of the species and the first record for Namibe Province. Our specimen was killed by locals when crossing a path early in the morning. The local name for this species is M’buta, which is also used for other large viperids such as the Puff-adder, Bitis arietans Merrem, 1820 (Ceria- co and Marques 2021).
Genus Causus Wagler, 1830
Causus nasalis Stejneger, 1893 Fig. 10c
Records. Mamué riparian area [-13.8006, 13.1230, 706 m] (CAS 263034; IN- BAC/AMB 10324).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 201
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
) “=. Woes © a . Figure 10. a Python natalensis ( errhis viperina f Lycophidion hellmichi g Psammophis subtaeniatus. Photographs by LMPC (a, ¢, f); AT (d, e, g) and DP (b).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 202
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Comments. Bocage (1895) suggested that Angolan populations of Causus resimus (Peters, 1862) belonged to a new variety, which he named angolensis. This decision was supported, according to the author, by morphological differ- ences between the Angolan populations and the nominotypical form. A few years earlier, Stejneger (1893) had already described what he called Causus nasalis from “West Africa” based on a specimen collected during the United States Eclipse Expedition to West Africa in 1890. Since Stejneger did not know the exact collection locality of the type, he assumed that it was “Cunga’” [most likely Fazenda Cunga, on the banks of the Kwanza River, Luanda Province]. A label bearing this information accompanied another specimen (USNM 16074), which we examined, and which is currently labelled as paratype in the collec- tions of the National Museum of Natural History, Smithsonian Institution, USA (USNM). Neither nasalis nor angolensis has been commented upon by subse- quent authors. The recently collected specimens from Serra da Neve could rep- resent this distinct Angolan population, considering not only its morphological differences but also its geographic isolation from the rest of the known distri- bution of the topotypical form. Our specimens agree entirely with the morpho- logical description provided by both Stejneger (1893) and Bocage (1895). Given this, we here recognize C. nasalis as a valid species for Angola, endemic to the coastal areas of the country, from Luanda to Namibe provinces.
Family Lamprophiidae Fitzinger, 1843 Genus Boaedon Dumeril, Bibron & Dumeéril. 1854
Boaedon variegatus Bocage, 1867 Fig. 10d
Records. vic. N’Dolondolo [-13.8105, 13.1361, 707 m] (CAS 263027); Catchi, basecamp [-13.7627, 13.2562, 1597 m] (MUNHAC/MB03-001536).
Comments. This species is a southwestern African endemic, distributed from the coastal areas of Kwanza Sul Province in Angola southwards to Na- mibia (Hallermann et al. 2020).
Genus Hemirhagerrhis Boettger, 1896
Hemirhagerrhis viperina (Bocage, 1873) Fig. 10e
Records. Basecamp 1 [-13.7770, 13.2591, 1488 m] (UF 187211); vic. N’Do- londolo [-13.8105, 13.1361, 291 m] (CAS 263028), [-13.8109, 13.1351, 705 ml] (CAS 263032); N’Dolondolo [-13.8133, 13.1362, 681 m] (INBAC/AMB 10279, 10298); Catchi, rock outcrops near basecamp [-13.7653, 13.2571, 1645 ml (MUNHAC/MB03-001537, 001538), [-13.7659, 13.2582, 1671 m] (MUNHAC/ MB03-001539); Catchi, basecamp [-13.7627, 13.2562, 1597 m] (MUNHAC/ MB03-001540, 001541).
Comments. This species is restricted to southwestern Angola and northern Namibia (Marques et al. 2018). It is relatively common across its range, where it is usually found during the day in rocky outcrops.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 203
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Genus Lycophidion Fitzinger, 1843
Lycophidion hellmichi Laurent, 1964 Fig. 10f
Records. Mauué riparian area [-13.8006, 13.1230, 706 m] (CAS 263033); Catchi surroundings [-13.7620, 13.2569, 1585 m] (MUNHAC/MB03-001 542, 001543). Comments. This recently collected material represents the first Angolan specimens of this species collected since its original description by Laurent (1964). The species appears to be endemic to southwestern Angola, from Ben- guela to Namibe Province and to northwestern Namibia. Our specimen from Serra da Neve was collected in the middle of its currently known distribution (Marques et al. 2018). A specimen from northwestern Namibia was assigned to this species by Broadley (1991, 1996). A very similar species, L. namibianum Broadley, 1991, which occurs in northwestern Namibia and southern Namibe Province (Broadley 1991, 1996; Lobon-Rovira et al. 2022), is distinguished from L. hellmichi by the lack of contact of the post-nasal with the first labial (in con- tact in L. hellmichi). Our specimens agree with the holotype in both coloration and the contact between post-nasal and first supralabial (Laurent 1964). The specimens from Catchi are adult male and female collected while copulating.
Genus Psammophis
Psammophis subtaeniatus Peters, 1882 Fig. 10g
Records. Maylowe [-13.8342, 13.2767, 803 m] (CAS 266140); Catchi surround- ings [-13.7620, 13.2569, 1585 m] (MUNHAC/MB03-001544); 2 km N of May- lowe [-13.8280, 13.2625, 803 m] (MUNHAC/MB03-001545).
Comments. Species restricted to dry shrublands and savannas, particularly Mopane woodland, and widely distributed from southern Angola and northern Namibia eastwards through Botswana, Zambia and Zimbabwe to western Mo- Zambique (Marques et al. 2018). Specimen MUNHAC/MB03-001545 was col- lected on sandy substrate near a rocky outcrop while active at night.
Genus Psammophylax Fitzinger, 1843
Psammophylax tritaeniatus (Giinther, 1868) Fig. 11a
Record. Catchi surroundings [-13.7620, 13.2569, 1585 m] (MUNHAC/MB03- 001546).
Comments. This species occurs in mid to high-elevation areas throughout southern and eastern Africa (Spawls et al. 2018). Previous records from Angola are from high-elevation areas of the Escarpment (Marques et al. 2018). This specimen represents the first record for Namibe Province and the westernmost record for P tritaeniatus. Further sampling across the species range may reveal additional cryptic diversity (Keates et al. 2019).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 204
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Family Colubridae Genus Dasypeltis Wagler, 1830
Dasypeltis scabra (Linnaeus, 1758) Fig. 11b
Record. N’Dolondolo [-13.8133, 13.1362, 681 m] (CAS 263023).
Comments. Following the key provided by Bates (2023), our specimen com- pletely conforms with Dasypeltis scabra. Marques et al. (2018) provided a map of the distribution of this widespread species in Angola.
Genus Telescopus Wagler, 1830
Telescopus semiannulatus polystictus Mertens, 1954 Fig. 11¢c
Records. vic. N’Dolondolo [-13.8105, 13.1361, 707 m] (CAS 263029); 2 km N of Maylowe [-13.8280, 13.2625, 803 m] (MUNHAC/MB03-001547).
Comments. The collected specimens exhibit more than 60 dark blotches on the back. This fits in the range attributed to the subspecies polystictus rather than to the nominotypical form (20-50 blotches according to Branch 1998). This is the first confirmed record for Angola of this subspecies that is otherwise known from neighboring Namibia.
Ne arte e at Eo ge Spe Pag” a s = “ “ - be Pe = oe : et ‘ por ni ’ 2 i lhl sae nP e _
Figure 11. a Psammophylax tritaeniatus b Dasypeltis scabra c Telescopus semiannulatus polystictus. Photographs by DP (a) and LMPC (b, c).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 205
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Discussion
Serra da Neve is situated in one of the most herpetologically rich areas not only in Angola, but in Southwest Africa more broadly (Marques et al. 2018). Due to its close proximity to the high elevation areas of the Angolan Escarpment, in combination with the surrounding lowland xeric areas of Namibe and Benguela provinces, the inselberg combines two distinct faunas: one associated with the Angolan high- lands (with some Zambezian or Congolian affinities), and one with the desert ar- eas extending from the Namib desert in Namibia to Benguela Province in Angola.
Despite its small area, the number of species recorded from the Serra da Neve inselberg is high, especially when strictly endemic species are considered. The number of Serra da Neve endemics is almost the same as in other biodiver- sity hotspots in the region, such as the Angolan Central Plateau and the Huila Escarpment, whose areas are many times the size of the inselberg. These num- bers are also considerably higher than those found on other similar inselbergs in Namibia. Whether these differences are related to the geomorphological and biogeographic characteristics of Serra da Neve, isolation, or simply reflect a bias due to poor sampling in other inselbergs, remains to be tested. Despite the already high number of recorded herpetological taxa for Serra da Neve and surrounding areas, several other taxa are expected to occur in the area, given their habitat preferences and known distribution ranges (Marques et al. 2018). Although amphibian diversity is lower in more arid than in mesic areas, sever- al other xeric-tolerant species, such as Poyntonophrynus dombensis (Bocage, 1895) and Sclerophrys regularis (Reuss, 1833), are expected to occur in Serra da Neve and nearby lowlands. The lack of records of members of the family Hyperoliidae, such as representatives of the Hyperolius angolensis Ahl, 1931 or H. benguellensis/nasutus species complexes, may be because none of our sampling events occurred in the peak of the rainy season. The habitats in the higher areas of the inselberg are similar to those where these species are found in other parts of the country. Other common rocket-frogs, such as Ptychadena oxyrhynchus (Smith, 1849) may also be expected in the study area. The same applies to certain squamate groups, such as representatives of the lacertid gen- era Ichnotropis and Nucras, amphisbaenids, rock monitors (Varanus albigularis angolensis Schmidt, 1933), or tree agamas of the genus Acanthocercus. To our surprise, despite the relatively high diversity of geckos in the area, we did not re- cord the ubiquitous Hemidactylus mabouia (Moreau de Jonnés, 1818), a species commonly found around human settlements. This may be another sign of the isolation from routes of regular human movements on and around the inselberg. Other species of geckos, such as the recently described Kolekanos spinicaudus Lobon-Rovira, Conradie, Baptista & Pinto, 2022, Rhoptropus benguellensis Mer- tens, 1938, and Pachydactylus cf. oreophilus McLachlan & Spence, 1967 may be present in the northern surrounding areas of the inselberg. Snake records always tend to be limited, especially during such short-term surveys and, there- fore, many other species are expected to occur in the area, such as Amblyodip- sas polylepis (Bocage, 1873), Aparallactus capensis Smith, 1849, Atractaspis bi- bronii Smith, 1849, Dispholidus typus (Smith, 1828), Prosymna sp., Philothamnus sp., Naja anchietae Bocage, 1879, and N. nigricincta Bogert, 1940 in both the lowland and highland areas; Bitis arietans, B. caudalis (Smith, 1839), and Aspi- delaps lubricus (Laurenti, 1768) in the lowlands; and Crotaphopeltis hotamboeia
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 206
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
(Laurenti, 1768), Dendroaspis polylepis Giinther, 1864, Elapsoidea sp., and Thel- otornis capensis oatesi (Giinther, 1881) in the highlands. The discovery of other unexpected and undescribed taxa cannot be ruled out, as seen from the recent descriptions of such species as Acontias mukwando and Lygodactylus baptistai.
The remoteness of Serra da Neve and the lack of good accesses to its base and summit have served so far as a guarantee of its preservation. Furthermore, the ruggedness, elevation, and climatic conditions of the inselberg may deter human settlement and concomitant large-scale habitat alterations. In ocean- ic islands, topographic and climatic conditions are known to have prevented land-cover changes, despite the presence of humans (Norder et al. 2020). This seems to apply to inselbergs such as Serra da Neve as well. A quick compari- son with Mount Moco, an inselberg of similar elevation and with similar number of human inhabitants seems to confirm this idea: Mount Moco is much less rugged than Serra da Neve, and much easier access by the local population has led to a considerable land-use change (Powell et al. 2023). Due to its remote- ness, Serra da Neve is unlikely to suffer from activities such as mining, that traditionally cause degradation of inselbergs (Porembski et al. 2016).
Landscape changes are clearly visible on the different plateaus of Serra da Neve, especially around human settlements with large agricultural fields and cattle pastures. Human-caused fires were observed during our fieldwork. The presence of livestock, such as cows, goats, and pigs were also noticeable, but no rodent species associated with humans, such as rats, were observed or collected in an ongoing small mammals survey. Traditional hunting is com- mon, targeting ungulates and birds, but not reptiles. However, chameleons and snakes are seen as dangerous and are usually killed when encountered, as it was the case of our specimens of Bitis gabonica, Telescopus semiannulatus polystictus, Psammphylax tritaeniatus, and Lycophidion hellmichi. At the base of the inselberg, tortoises, such as Kinixys, are consumed or traded as a delicacy. The remaining amphibians and reptiles are usually neglected by the local hu- man population. Pollution is noticeable in a few water bodies due to its use by humans and cattle and may pose a threat to some amphibians.
Serra da Neve was identified as a potential conservation area by both the scientific community (Huntley and Matos 1992; Huntley 2010; Pinto et al. 2023) and the Angolan authorities. The present checklist, even though it is focused only on amphibians and reptiles, unambiguously shows its conservation impor- tance and interest, not only due to its high number of endemic taxa, but also to its taxonomic diversity. Similar checklists are currently being prepared for other taxonomic groups, such as birds and mammals (Marks pers. comm. November 2022; Ferguson pers. comm. November 2022). This is particularly important in the current scenario of climate change, as inselbergs in xeric areas can serve as important biodiversity retreats (Burke 2003).
Acknowledgements
The present work is a result of the ongoing collaboration between the Insti- tuto Nacional de Biodiversidade e Areas de Conservacao (INBAC) from the Ministry of Environment of Angola and its international partners. Angolan specimens were collected and exported under permits issued by INBAC (155/ INBAC.MINAB/2017; 28/INBAC.MINAMB/2019; 65/INBAC.MINAMB/2022).
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 207
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
We also thank the provincial and local authorities for their support and coop- eration during our fieldwork. We thank Ishan Agarwal, Joyce Janota, Suzana Bandeira, Adam Ferguson, Ben Marks and Daryl Coldren for their support during field work. Special thanks to Alvaro (“Varito”) Baptista and his team from Omauha Lodge, for all the assistance, great support, and friendship during the field work.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
This work was funded by the National Geographic Society Explorer Grant (NGS- 73084R-20) to LMPC. Fundacao para a Ciéncia e Tecnologia (FCT) supported MPM un- der grants (SFRH/BD/129924/2017, COVID/BD/152155/2022), DP under grant (2021. 05238.BD), and MLL under contract (2020. 03608.CEECIND). Work supported by the European Union's Horizon 2020 Research and Innovation Programme under the Grant Agreement Number 857251. AMB and LMPC were supported by grant DEB 2146654 from the National Science Foundation of the United States.
Author contributions
Conceptualization: MPM, LMPC. Data curation: MPM. Formal analysis: MPM, DP, MLL, AMB, AT, LMPC. Funding acquisition: AMB, LMPC. Investigation: MPM, DP LMPC, AT, AMB. Methodology: MPM, DP, MLL, LMPC. Project administration: LMPC, AMB. Soft- ware: MLL. Supervision: LMPC, AMB. Writing — original draft: MPM, DP. Writing — review and editing: AMB, MLL, LMPC, AT.
Author ORCIDs
Mariana P. Marques © https://orcid.org/0000-0002-1 712-2632 Diogo Parrinha © https://orcid.org/0000-0002-1302-025xX Manuel Lopes-Lima © https://orcid.org/0000-0002-2761-7962 Arthur Tiutenko © https://orcid.org/0000-0001-5952-4923 Aaron M. Bauer ® https://orcid.org/0000-0001-6839-8025 Luis M. P. Ceriaco © https://orcid.org/0000-0002-0591-9978
Data availability
All of the data that support the findings of this study are available in the main text.
References
Adalsteinsson SA, Branch WR, Trape S, Vitt LU, Hedges SB (2009) Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squamata). Zootaxa 2244(1): 1-50. https://doi.org/10.11646/zootaxa.2244.1.1
Baptista NL, Pinto PV, Keates C, Lobon-Rovira J, Edwards S, Rodel M-O (2023) Two new Poyntonophrynus species (Anura: Bufonidae) highlight the importance of Angolan
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 208
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
centers of endemism. Vertebrate Zoology 73: 991-1031. https://doi.org/10.3897/ vz./3.e103935
Barker N, Clarck R, Neef G, Maiato F, Francisco D, Goyder D (2015) Learning the ABCs: Angolan botanical collecting. Part 2: bush pilots and old volcanoes. Veld and Flora 101(3): 116-119.
Barlow A, Wister W, Kelly CMR, Phelps T, Tolley KA (2019) Ancient habitat shifts and organismal diversification are decoupled in the African viper genus Bitis (Serpentes: Viperidae). Journal of Biogeography 2019: 1-15. https://doi.org/10.1111/jbi.13578
Bates MF (2023) The enigmatic Namibian egg-eating snake Dasypeltis loveridgei Mer- tens, 1954 (Squamata: Colubridae): A valid species recovered from synonymy. Bonn Zoological Bulletin 72(2): 163-184. https://doi.org/10.20363/BZB-2023.72.2.163
Bauer AM, Lamb T (2005) Phylogenetic relationships of southern African geckos in the Pachydactylus group (Squamata: Gekkonidae). African Journal of Herpetology 54: 105-129. https://doi.org/10.1080/21564574.2005.9635525
Bauer AM, Sadlier RA, Jackman TR (2022) A revision of the genus Bavayia Roux, 1913 (Squamata: Gekkota: Diplodactylidae), a non-adaptive radiation of microendemic species. Proceedings of the California of Sciences 67(Supplement |): 1-236.
Bauer AM, Ceriaco LMP Marques MP, Becker FS (2023) Highland reptiles of Angola and Namibia. Namibian Journal of Environment 8: 259-276.
Bayliss J, Timberlake J, Branch WR, Bruessow C, Collins S, Congdon C, Curran M, de Sousa C, Dowsett R, Dowsett-Lemaire F, Fishpool L, Harris T, Herrmann E, Georgiadis S, Kopp M, Liggit B, Monadjem A, Patel H, Ribeiro D, Spottiswoode C, Taylor P Willcock S, Smith P (2014) The discovery, biodiversity and conservation of Mabu forest — the largest medium-altitude rainforest in southern Africa. Oryx 48(2): 177-185. https:// doi.org/10.101 7/S0030605313000720
Becker FS, Baptista NL, Pinto PV, Ernst R, Conradie W (2023) The amphibians of the high- lands and escarpments of Angola and Namibia. Namibian Journal of Environment 8: 245-257.
Bilton DT (2014) Two new water beetles from the Hantamsberg, an inselberg in the Northern Cape of South Africa (Coleoptera, Hydraenidae). Zootaxa 3887(4): 471- A480. https://doi.org/10.11646/zootaxa.3887.4.5
Bittencourt-Silva, GB, Bayliss J, Conradie W (2020) First herpetological surveys of Mount Lico and Mount Socone, Mozambique. Amphibian and Reptile Conservation 14(2): 198-217 (e247).
Bocage JVB (1895) Herpétologie d’Angola et du Congo. Imprimerie Nationale, Lisbon, 203 pp. [+ 20 pls]
Boulenger GA (1907) Descriptions of three new lizards and a new frog, discovered by Dr. W.J. Ansorge in Angola. Annals & Magazine of Natural History 7(19): 212-214. https://doi.org/10.1080/00222930709487258
Branch WR (1998) Field Guide to the Snakes and other Reptiles of Southern Africa (3 ed.). Struik Publisher, Cape Town, 399 pp.
Branch WR, Rédel M-O, Marais J (2005) A new species of rupicolous Cordylus Laurenti 1768 (Sauria: Cordylidae) from Northern Mozambique. African Journal of Herpetolo- gy 131-138. https://doi.org/10.1080/21 564574.2005.9635526
Branch WR, Bayliss J (2009) A new species of Atheris (Serpentes: Viperidae) from northern Mozambique. Zootaxa 2113(1): 41-54. https://doi.org/10.11646/zootaxa.2113.1.2
Branch WR, Tolley KA (2010) A new species of chameleon (Sauria: Chamaeleonidae: Nadzikambia) from Mount Mabu, central Mozambique. African Journal of Herpetolo- gy 59(2): 157-172. https://doi.org/10.1080/21564574.2010.516275
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 209
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Branch WR, Bayliss J, Tolley KA (2014) Pygmy chameleons of the Rhampholeon platy- ceps complex (Squamata: Chamaeleonidae): Description of four new species from isolated ‘sky islands’ of northern Mozambique. Zootaxa 3718(1): 1-36. https://doi. org/10.11646/zootaxa.3814.1.1
Branch WR, Bayliss K, Bittencourt-Silva GB, Conradie W, Engelbrecht HM, Loader SP Me- negon M, Nanvonamuduitxo C, Tolley KA (2019) A new species of tree snake (Dipsa- doboa, Serpentes: Colubridae) from ‘sky island’ forests in northern Mozambique, with notes on other members of the Dipsodoboa werneri group. Zootaxa 4646(3): 541-563. https://doi.org/10.11646/zootaxa.4646.3.6
Branch WR, Schmitz A, Lobon-Rovira J, Baptista NL, Antonio T, Conradie W (2021) Rock Island Melody: A revision of the Afroedura bogerti Loveridge, 1944 group, with de- scriptions of four new endemic species from Angola. Zoosystematics and Evolution 97(1): 55-82. https://doi.org/10.3897/zse.97.57202
Brand RF, Brown LR, Du Preez PJ (2011) Biogeography of Platberg, Eastern Free State, South Africa: Links with Afromontane Regions and South African Biomes. In: Grillo O, Venora G (Eds) Biodiversity Loss in a Changing Planet. InTech Publisher, Internation- al, 113-136. https://doi.org/10.5772/24916
Brand RF, Scott-Shaw CR, O’Connor TG (2019) The alpine flora on inselberg summits in the Maloti-Drakensberg Park, KwaZulu-Natal, South Africa. Bothalia 49(1): a2386. https://doi.org/10.4102/abc.v49i1.2386
Broadley DG (1991) A review of the Namibian snakes of the genus Lycophidion (Ser- pentes: Colubridae), with the description of a new endemic species. Annals of the Transvaal Museum 35(14): 209-215.
Broadley DG (1996) A revision of the genus Lycophidion Fitzinger (Serpentes: Colubri- dae) in Africa South of the Equator. Syntarsus 3: 1-33.
Broadley DG, Broadley S (1999) A review of the African Worm Snakes from south of latitude 12°S (Serpentes: Leptotyphlopidae). Syntarsus 5: 1-36.
Burke A (2001) Determinants of inselberg floras in arid Nama Karoo landscapes. Journal of Biogeography 28(10): 1211-1220. https://doi.org/10.1046/j.1365- 2699.2001.00623.x
Burke A (2003) The role of Namibian inselbergs in contributing to local and regional plant species richness. Biodiversity and Conservation 12(3): 469-486. https://doi. org/10.1023/A:1022491 723262
Burke A (2005) Biodiversity Patterns in Arid, Variable Environments A case study of Na- mibian inselberg and mountain flora. Mountain Research and Development 25(3): 228-234. https://doi.org/10.1659/0276-4741(2005)025[0228:BPIAVE]2.0.CO;2
Butler BO, Ceriaco LMP Marques MP, Bandeira S, Julio T, Heinicke M, Bauer AM (2019) Herpetological survey of Huila Province, Southwest Angola, including first records from Bicuar National Park. Herpetological Review 50(2): 225-240.
Butler BO, Ceriaco LMP Jackman TR, Bauer AM (in press) Population genetics and phy- logeopgraphy of Trachylepis sulcata (Peters, 1862) and T. ansorgii (Boulenger, 1907) in southwestern Africa. African Journal of Herpetology.
Ceriaco LMP Marques MP (2021) Serpentes Venenosas de Angola. Guia de Identifi- cacao e Primeiros Socorros. Editora Arte & Ciéncia, Porto, 216 pp.
Ceriaco LMP de Sa SAC, Bandeira S, Valério H, Stanley EL, Kuhn AL, Marques MP Vin- dum JV, Blackburn DC, Bauer AM (2016) Herpetological survey of lona National Park and Namibe Regional Natural Park, with a synoptic list of the amphibians and reptiles of Namibe Province, southwestern Angola. Proceedings of the California Academy of Sciences 63(2): 15-61.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 210
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Ceriaco LMP, Marques MP Bandeira S, Agarwal |, Stanley EL, Bauer AM, Heinicke MP, Blackburn DC (2018) A new earless species of Poyntonophrynus (Anura, Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola. ZooKeys 780: 109-136. https://doi.org/10.3897/zookeys.780.25859
Ceriaco LMP, Tolley KA, Marques MP, Heinicke MP, Bauer AM (2020a) A dwarf among gi- ants: Phylogenetic position of the elusive Angolan Adder (Bitis heraldica) and biogeo- graphic affinities of Angolan Afromontane regions. African Journal of Herpetology 69(1): 108-118. https://doi.org/10.1080/21564574.2020.1782484
Ceriaco LMP, Heinicke MP, Parker KL, Marques MP Bauer AM (2020b) A review of the Af- rican snake-eyed skinks (Scincidae: Panaspis) from Angola, with the description of a new species. Zootaxa 4747(1): 77-112. https://doi.org/10.11646/zootaxa.4747.1.3
Ceriaco LMP, Agarwal |, Marques MP, Bauer AM (2020c) A review of the genus Hemidac- tylus Goldfuss, 1820 (Squamata: Gekkonidae) from Angola, with the description of two new species. Zootaxa 4746(1): 1-71. https://doi.org/10.11646/zootaxa.4746.1.1
Ceriaco LMP Santos B, Marques MP, Bauer AM, Tiutenko A (2021) Citizen Science medi- as meets specimens in old formalin filled jars: A new species of Banded Rubber Frog, genus Phrynomantis (Anura: Mycrohylidae) from Angola. Alytes 38(1—4): 18-48.
Ceriaco LMP, Marques MP Bell RC, Bauer AM (2022) The terrestrial reptiles of the Gulf of Guinea Oceanic Islands. In: Cerfaco LMP, Lima RF, Melo M, Bell RC (Eds) Biodiversity of the Gulf of Guinea Oceanic Islands. Science and Conservation. Springer Nature, Cham, 505-534. https://doi.org/10.1007/978-3-031-06153-0_19
Ceriaco LMP, Marques MP Parrinha D, Tiutenko A, Weinell J, Butler BO, Bauer AM (2024) The Trachylepis (Squamata: Scincidae) of Angola: an integrative taxonomic review with the description of seven new species. Bulletin of the American Museum of Natu- ral History 465(465): 1-153. https://doi.org/10.1206/0003-0090.465.1.1
Channing A (2001) Amphibians of Central and Southern Africa. Cornell University Press, Ithaca, New York, 470 pp. https://doi.org/10.7591/9781501 733697
Channing A, Baptista N (2013) Amietia angolensis and A. fuscigula (Anura: Pyxicephali- dae) in southern Africa: A cold case reheated. Zootaxa 3640(4): 501-520. https://doi. org/10.11646/zootaxa.3640.4.1
Channing A, Becker F (2019) Correction to the type locality of Tomopterna ahli (Deckert, 1938) (Anura: Pyxicephalidae), with the designation of a neotype. Zootaxa 4688(4): 549-560. https://doi.org/10.11646/zootaxa.4688.4.6
Channing A, Rédel M-O (2019) Field guide to the frogs and other amphibians of Africa. Struik Nature, Cape Town, 408 pp.
Channing A, Dehling JM, Létters S, Ernst R (2016) Species boundaries and taxonomy of the African river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa 4155(1): 1-76. https://doi.org/10.11646/zootaxa.41 55.1.1
Colston TJ, Pyron AR, Bauer AM (2020) A new species of African snake-eyed skink (Scin- cidae: Panaspis) from Ethiopia. Zootaxa 4779(2): 190-200. https://doi.org/10.11646/ zootaxa.4/79.2.2
Congdon TCE, Collins S, Bayliss J (2010) Butterflies of south-east Africa’s mountains (Mozambique and Malawi). Metamorphosis 21(2): 45-107.
Conradie W, Measey GJ, Branch WR, Tolley KA (2012) Revised phylogeny of African sand lizards (Pedioplanis), with the description of two new species from south-western Angola. African Journal of Herpetology 61(2): 91-112. https://doi.org/10.1080/215 64574.2012.676079
Conradie W, Bittencourt-Silva G, Engelbrecht HM, Loader SR Menegon M, Nan- vonamuduitxo C, Scott M, Tolley KA (2016a) Exploration into the hidden world of Mo-
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 211
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
zambique's sky island forests: New discoveries of reptiles and amphibians. Zoosys- tematics and Evolution 92(2): 163-180. https://doi.org/10.3897/zse.92.9948
Conradie W, Bills R, Branch WR (2016b) The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola. Amphibian and Reptile Con- servation 10(2)[Special Section]: 6-36.
Conradie W, Bittencourt-Silva GB, Faroog HM, Loader SP. Menegon M, Tolley KA (2018) New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozam- bique inselbergs. African Journal of Herpetology 61-85. https://doi.org/10.1080/21 564574.2017.1376714
Conradie W, Keates C, Verburght L, Baptista NL, Harvey J, Julio T, Neef G (2022) Contri- butions to the herpetofauna of the Angolan Okavango-Cuando-Zambezi River drain- ages. Part 2: Lizards (Sauria), chelonians, and crocodiles. Amphibian and Reptile Conservation 16(2): 181-214 (e322).
Conradie W, Lobon-Rovira J, Becker GS, Schmitz A, Pinto PV (2023) Flat gecko (Afroed- ura) diversity, endemism and speciation in the highlands and escarpments of Angola and Namibia. Namibian Journal of Environment 8: 277-281.
Curran M, Koop M, Beck J, Fahr J (2012) Species diversity of bats along an altitude gra- dient on Mount Mulanje, southern Malawi. Journal of Tropical Ecology 28: 243-253. https://doi.org/10.1017/S026646741 2000193
Daniels SR, Bayliss J (2012) Neglected refugia of biodiversity: mountainous regions in Mozambique and Malawi yield two novel freshwater crab species (Potamonautidae: Potamonautes). Zoological Journal of the Linnean Society 164(3): 498-509. https:// doi.org/10.1111/j.1096-3642.2011.00773.x
Daniels S, Phiri E, Bayliss J (2014) Renewed sampling of inland aquatic habitats in southern Africa yields two novel freshwater crab species (Decapoda: Potamonau- tidae: Potamonautes). Zoological Journal of the Linnean Society 171(2): 356-369. https://doi.org/10.1111/zoj.12139
Daniels SR, Bittencourt-Silva GB, Muianga V, Bayliss J (2020) Phylogenetics of the fresh- water crab (Potamonautes MacLeay, 1838) fauna from ‘sky islands’ in Mozambique with the description of a new species (Brachyura: Potamoidea: Potamonautidae). Eu- ropean Journal of Taxonomy 716(716): 1-23. https://doi.org/10.5852/ejt.2020.716
Elzen P van den (1983) Zur Herpetofauna des Brandberges. Bonner zoologische Beit- rage 34: 293-309.
Fouquet A, Kok PJR, Recoder RS, Prates |, Camacho A, Marques-Souza S, Ghellere JM, McDiarmid RW, Rodrigues MT (2023) Relicts in the mist: Two new frog families, genera and species highlight the role of Pantepui as a biodiversity museum through- out the Cenozoic. Molecular Phylogenetics and Evolution 191: 107971. https://doi. org/10.1016/j.ympev.2023.107971
Frétey T, Dewynter M, Ohler A (2018) Onymotopes in zoological nomenclature: Some addi- tional terms, with fixation of a lectonymotope for Xenopus petersii Bocage, 1895 (Am- phibia, Anura). Bionomina 13(1): 37-50. https://doi.org/10.11646/bionomina.13.1.3
Fritz U, Tolley KA, Vamberger M, Ihlow F (2022) Phylogeny and phylogeography of chelo- nians from sub-Saharan Africa — A review of current knowledge in tribute to Margaretha D. Hofmeyr. Vertebrate Zoology 72: 951-969. https://doi.org/10.3897/vz.72.e95681
Furman BLS, Bewick AJ, Harrison TL, Greenbaum E, Gvozdik V, Kusamba C, Evans BJ (2015) Pan-African phylogeography of a model organism, the African clawed frog Xen- opus laevis. Molecular Ecology 24(4): 909-925. https://doi.org/10.1111/mec.13076
Goudie A, Viles H (2015) Landscapes and Landforms of Namibia. Springer, Dordrecht, 173 pp. https://doi.org/10.1007/978-94-01 7-8020-9
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 212
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Grandvaux-Barbosa LA (1970) Carta fitogeografica de Angola. Instituto de Investigacado Cientifica de Angola, Luanda, 323 pp.
Griffin M (2000) Annotated checklist of amphibians, reptiles and mammals of the Brand- berg, central Namib Desert, Namibia. Cimbebasia Memoir 9: 69-89.
Hallermann J, Ceriaco LMP Schmitz A, Ernst R, Conradie W, Verburgt L, Marques MP Bauer AM (2020) A review of the Angolan House snakes, genus Boaedon Duméril, Bibron and Dumeéril (1854) (Serpentes: Lamprophiidae), with description of three new species in the Boaedon fuliginosus (Boie, 1827) species complex. African Journal of Herpetology 69(1): 29-78. https://doi.org/10.1080/21564574.2020.1777470
Heinicke MP, Adderly LM, Bauer AM, Jackman TR (2011) A long-known new species of gecko allied to Pachydactylus bicolor (Squamata: Gekkonidae) from central Namib- ian coast. African Journal of Herpetology 60: 113-129. https://doi.org/10.1080/215 64574.2011.613952
Heinicke MP, Ceriaco LMP, Moore IM, Bauer AM, Blackburn DC (2017a) Tomopterna damarensis (Anura: Pyxicephalidae) is broadly distributed in Namibia and Angola. Salamandra (Frankfurt) 53(3): 461-465.
Heinicke MP Jackman TR, Bauer AM (2017b) The measure of success: Geographic iso- lation promotes diversification in Pachydactylus geckos. BMC Evolutionary Biology 17(9): 1-17. https://doi.org/10.1186/s12862-016-0846-2
Heinz HM (2011) Comparative phylogeography of two widespread geckos from typically narrow-ranging Pachydactylus group in southern Africa. Unpublished MS thesis. Villa- nova University, Pennsylvania.
Heinz MD, Brennan IG, Jackman TR, Bauer AM (2021) Phylogeny of the genus Chon- drodactylus (Squamata: Gekkonidae) with the establishment of a stable taxono- my. Bulletin of the Museum of Comparative Zoology 163(5): 151-210. https://doi. org/10.3099/0027-4100-163.5.151
Hellmich W (1957a) Herpetologische Ergebnisse einer Forschungsreise in Angola. Veroffentlichungen der Zoologischen Staatssammlung Munchen 5: 1-92.
Hellmich W (1957b) Die Reptilienausbeute der hamburgischen Angola-Expedition. Mit- teilungen aus dem Hamburgischen Zoologischen Museum und Institut 55: 39-80. Huntley BJ (2010) Estratégia de expansdao da rede de areas protegidas de Angola/Pro- posals for na Angolan protected area expansion strategy (APAES). Unpublished re-
port: Ministry of Environment, Luanda.
Huntley BJ (2023) Biomes and ecoregions of the highlands and escarpment of Angola and Namibia. Namibian Journal of Environment 8: 29-41.
Huntley BJ, Matos EM (1992) Biodiversity: Angolan environmental status quo assess- ment report. Unpublished report: IUCN Regional Office for Southern Africa, Harare, Zimbabwe, 55 pp.
Kandziora M, Gehrke B, Popp M, Gizaw A, Brochmann C, Pirie MD (2022) The enigmatic tropical alpine flora on the African sky islands is young, disturbed, and unsaturated. Proceedings of the National Academy of Sciences of the United States of America 119(22): e2112737119. https://doi.org/10.1073/pnas.2112737119
Keates C, Conradie W, Greenbaum E, Edwards S (2019) A snake in the grass: Genetic struc- turing of the widespread African grass snake (Psammophylax Fitzinger, 1843), with the description of anew genus and a new species. Journal of Zoological Systematics and Evolutionary Research 57(4): 1039-1066. https://doi.org/10.1111/jzs.12337
Kirchhof S, Kramer M, Linden J, Richter K (2010) The reptile species assemblage of the Soutpansberg (Limpopo Province, South Africa) and its characteristics. Salamandra (Frankfurt) 46(3): 147-166.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 213
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Kuhn AL (2016) Systematics of the Namib day geckos (Squamata: Gekkonidae: Rhoptro- pus). Unpublished MS thesis. Villanova University, Pennsylvania.
Laurent RF (1964) Reptiles et amphibiens de I'Angola (Troisieme contribution). Publi- cacoes Culturais da Companhia de Diamantes de Angola 67: 1-165.
Lob6n-Rovira J, Conradie W, Iglesis DB, Ernst R, Verissimo L, Baptista N, Pinto PV (2021) Between sand, rock and branches: An integrative taxonomic revision of Angolan Hemidactylus Goldfuss, 1820, with description of four new species. Vertebrate Zool- ogy 71: 465-501. https://doi.org/10.3897/vz.71.e64781
Lobon-Rovira J, Pinto PV, Becker FS, Tolley KA, Measey J, Boon B, de Sa S, Conradie W (2022) An updated herpetofaunal species inventory of lona National Park in south- western Angola. Check List 18(2): 289-321. https://doi.org/10.15560/18.2.289
MacArthur RH, Wilson EO (2001) The Theory of Island Biogeography. Princeton Universi- ty Press, Princeton, 203 pp. https://doi.org/10.1515/9781400881376
Main DC, van Vuuren BJ, Tilbury CR, Tolley KA (2022) Out of southern Africa: Origins and cryptic speciation in Chamaeleo, the most widespread chameleon genus. Molecular Phy- logenetics and Evolution 175: 107578. https://doi.org/10.1016/j.ympev.2022.107578
Marques MP, Ceriaco LMP, Blackburn DC, Bauer AM (2018) Diversity and Distribution of the Amphibians and Terrestrial Reptiles of Angola - Atlas of Historical and Bib- liographic Records (1840-2017). Proceedings of the California Academy of Sciences 4(65, Supplement II): 1-501.
Marques MP Ceriaco LMP Stanley EL, Bandeira AS, Agarwal |, Bauer AM (2019) A new species of girdled lizard (Squamata: Cordylidae) from the Serra da Neve Inselberg, Namibe Province, southwestern Angola. Zootaxa 4668(4): 503-524. https://doi. org/10.11646/zootaxa.4668.4.4
Marques MP, Ceriaco LMP Buehler MD, Bandeira SA, Janota JM, Bauer AM (2020) A revi- sion of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with the description of three new species. Zootaxa 4853(3): 301-352. https://doi. org/10.11646/zootaxa.4853.3.1
Marques MP Ceriaco LMP Heinicke MP Chehouri RM, Conradie W, Tolley KA, Bauer AM (2022) The Angolan bushveld lizards, genus Heliobolus Fitzinger, 1843 (Squamata: Lacertidae): Integrative taxonomy and the description of two new species. Vertebrate Zoology 72: 745-769. https://doi.org/10.3897/vz.72.e85269
Marques MP Parrinha D, Ceriaco LMP, Brennan IG, Heinicke MP, Bauer AM (2023a) A new species of thick-toed gecko (Pachydactylus) from Serra da Neve and surrounding rocky areas of southwestern Angola. Vertebrate Zoology 73: 325-343. https://doi. org/10.3897/vz.73.e101329
Marques MP, Parrinha D, Tiutenko A, Lopes-Lima M, Bauer AM, Ceriaco LMP (2023b) A new species of African legless-skink, genus Acontias Cuvier, 1818 “1817” (Squamata: Scincidae) from Serra da Neve inselberg, southwestern Angola. African Journal of Herpetology 7(2): 145-162. https://doi.org/10.1080/21564574.2023.2246487
Medina MF, Bauer AM, Branch WR, Schmitz A, Conradie W, Nagy ZT, Hibbitts TJ, Ernst R, Portik DM, Nielsen SV, Colston TJ, Kusamba C, Behangana M, Rodel M-O, Greenbaum E (2016) Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evo- lution 100: 409-423. https://doi.org/10.1016/j.ympev.2016.04.026
Mendelsohn JM, Huntley BJ (2023) Introducing the highlands and escarpments of An- gola and Namibia. Namibian Journal of Environment 8: 7-22.
Mertens R (1938) Amphibien und Reptilien aus Angola gesammelt von W. Schack. Seck- enbergiana 20(6): 425-443.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 214
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Michael DR, Cunningham RB, Lindenmayer DB (2008) A forgotten habitat? Granite insel- bergs conserve reptile diversity in fragmented agricultural landscapes. Journal of Ap- plied Ecology 45(6): 1742-1752. https://doi.org/10.1111/j.1365-2664.2008.01567.x
Monadjem A, Schoeman MC, Reside A, Pio DV, Stoffberg S, Bayliss J, Cotterill FPD, Cur- ran M, Kopp M, Taylor PJ (2010) A recent inventory of the bats of Mozambique with documentation of seven new species for the country. Acta Chiropterologica 12: 371- 391. https://doi.org/10.3161/150811010X537963
Monard A (1937) Contribution a I'herpétologie d'Angola. Arquivos do Museu Bocage 8: 19-154.
Norder SJ, Lima RF, Nacimento L, Lim JY, Fernandez-Palacios JM, Romeiras MM, Elias RB, Cabezas FJ, Catarino L, Ceriaco LMP Castilla-Beltran A, Gabriel R, Sequeira MM, Rijsdijk KF, Nogué S, Kissling WD, van Loon EE, Hall M, Matos M, Borges PAV (2020) Global change in microcosms: Environmental and societal predictors of land cov- er change on the Atlantic Ocean Islands. Anthropocene 30: 100242. https://doi. org/10.1016/j.ancene.2020.100242
Parrinha D, Marques MP, Heinicke MP Khalid F, Parker KL, Tolley KA, Childers JL, Con- radie W, Bauer AM, Cerfaco LMP (2021) A revision of the genus Pedioplanis Fitzinger (Squamata: Lacertidae) from Angola, with the description of a new species. Zootaxa 5032: 1-46. https://doi.org/10.11646/zootaxa.5032.1.1
Pereira E (1977) Serra da Neve (Angola) — Nota sobre a geomorfologia da regido e idade das aplanacoes. Boletim da Sociedade Geologica de Portugal 20: 277-282.
Perret J-L (1976) Révisions des amphibiens africains et principalement des types, con- servés au Musée Bocage de Lisbonne. Arquivos do Museu Bocage 2(6): 15-34.
Pinto PV, Russo V, Verissimo L (2023) The highlands in Angolan conservation areas. Namibian Journal of Environment 8: 53-62.
Porembski S (2007) Tropical inselbergs: Habitat types, adaptive strategies and diversity patterns. Revista Brasileira de Botanica. Brazilian Journal of Botany 30(4): 579-586. https://doi.org/10.1590/S0100-84042007000400004
Porembski S, Barthlott W (2000) Granitic and gneissic outcrops (inselbergs) as cen- ters of diversity for desiccation-tolerant vascular plant. Plant Ecology 151(1): 19-28. https://doi.org/10.1023/A:102656581 7218
Porembski S, Silveira FAO, Fiedler PL, Watve A, Rabarimanarivo M, Kouame F, Hopper SD (2016) World destruction of inselbergs and related rock outcrops threatens a unique ecosystem. Biodiversity and Conservation 25(13): 2827-2830. https://doi. org/10.1007/s10531-016-11 71-1
Porembski S, Silveira FAO, Fiedler PL, Watve A, Rabarimanarivo AW, Kouame F, Hopper SD (2017) Worldwide destruction of inselbergs and related rock outcrops threatens a unique ecosystem. Biodiversity and Conservation 25(13): 2827-2839. https://doi. org/10.1007/s10531-016-1171-1
Portik DM, Mulungu E, Sequeira D, McEntee JP (2013a) Herpetological surveys of the Serra Jeci and Namuli massifs, Mozambique, and an annotated checklist of the Southern Afromontane Archipelago. Herpetological Review 44: 394-406.
Portik DM, Travers SL, Bauer AM, Branch WR (2013b) A new species of Lygodactylus (Squamata: Gekkonidae) endemic to Mt. Namuli, an isolated ‘sky island’ of northern Mozambique. Zootaxa 3710(5): 415-435. https://doi.org/10.11646/zootaxa.3710.5.2
Powell LL, Pinto PV, Mills MSL, Baptista NL, Costa K, Dijktra K-DB, Gomes AL, Guedes P Julio T, Monadjem A, Palmeirim AF, Russo V, Melo M (2023) The last Afromontane forests in Angola are threatened by fires. Nature Ecology & Evolution 7(5): 628-629. https://doi.org/10.1038/s41559-023-02025-9
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 215
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Poynton JC, Haacke WD (1993) On a collection of amphibians from Angola, including a new species of Bufo Laurenti. Annals of the Transvaal Museum 36: 1-16.
Poynton JC, Loader SP, Conradie W, Rodel M-O, Liedtke HC (2016) Designation and de- scription of a neotype of Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1837) (Amphibia: Anura: Bufonidae). Zootaxa 4098(1): 73-91. https://doi.org/10.11646/zootaxa.4098.1.3
Rabarimanarivo MN, Ramandimbisoa B, Rakotoarivelo NH, Phillipson PB, Andriambololonera S, Callmander MW, Porembski S (2019) The extraordinary botanical diversity of insel- bergs in Madagasar. Candollea 74(1): 65-84. https://doi.org/10.15553/c2019v741a8
Rapson SA, Goldizen AW, Seddon JM (2013) Gene flow in mongooses endemic to Namibia's granite inselbergs despite past climatic fluctuations and isolating land- scape features. Journal of Mammalogy 94(1): 218-230. https://doi.org/10.1644/11- MAMN-A-379.1
Recoder R, Prates I, Marques-Souza S, Camacho A, Nunes PMS, Vechio FD, Ghellere JM, McDiarmid RW, Trefaut Rodrigues M (2020) Lizards from the Lost World: Two new species and evolutionary relationships of the Pantepui highland Riolama (Gymnoph- thalmidae). Zoological Journal of the Linnean Society 190(1): 271-297. https://doi. org/10.1093/zoolinnean/zlz168
Roux-Estéve R (1974) Révision systématique des Typhlopidae d'Afrique. Reptilia. Ser- pentes. Memdires du Muséum National d'Histoire Naturelle 87: 1-313.
Schigtz A (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.
Schmidt KP (1933) The reptiles of the Pulitzer-Angola Expedition. Annals of the Carne- gie Museum 22: 1-15. https://doi.org/10.5962/p.214561
Simmons RE, Griffin M, Griffin RE, Marais E, Kolberg H (1998) Endemism in Namibia: Patterns, processes, and predictions. Biodiversity and Conservation 7(4): 513-530. https://doi.org/10.1023/A:1008879712736
Simmons NB, Flanders J, Fils EMB, Parker G, Suter JD, Bamba S, Douno M, Keita MK, Morales AE, Frick WF (2021) A new dichromatic species of Myotis (Chiroptera: Ver- pertilionidae) from the Nimba Mountains, Guiniea. American Museum Novitates 2963(3963): 1-37. https://doi.org/10.1206/3963.1
Spawls S, Howell K, Hinkel H, Menegon M (2018) Field Guide to East African Reptiles. Bloomsbury, London, 554 pp.
Spawls S, Mohammad A, Mazuch T (2023) Handbook of Amphibians and Reptiles of North-east Africa. Bloomsbury Publishing 1% edn., 640 pp.
Stejneger L (1893) On some collections of reptiles and batrachians from East Africa and the adjacent islands, recently received from Dr. W.L. Abbott and Mr. William Astor Chanler, with descriptions of new species. Proceedings of the United States National Museum 16(970): 711-741. https://doi.org/10.5479/si.00963801.970.711
Taylor PJ, Stoffberg S, Monadjem A, Schoeman MC, Bayliss J, Cotterill FPD (2012) Four new bat species (Rhinolophus hildebrandtii complex) reflect Plio-Pleistocene diver- gence of dwarfs and giants across an afromontane archipelago. PLOS ONE 7(9): e41744. https://doi.org/10.1371/journal.pone.0041 744
Tilbury C (2010) Chameleons of Africa: An Atlas, Including the Chameleons of Europe, the Middle East and Asia. Edition Chimaira, Frankfurt am Main, 831 pp.
Timberlake JR, Dowsett-Lemaire F, Bayliss J, Alves T, Baena S, Bento C, Cook K, Francis- co J, Harris T, Smith PB de Sousa C (2009) Mt. Namuli, Mozambique: Biodiversity and Conservation. Report produced under the Darwin Initiative Award 15/036: Biodiver- sity Loss in South-East Africa’s Montane Ecosystems. Royal Botanic Gardens, Kew, London, 114 pp.
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 216
Mariana P. Marques et al.: Herpetofauna of the Serra da Neve inselberg, Southwestern Angola
Weinell JL, Bauer AM (2018) Systematics and phylogeography of the widely distributed African skink Trachylepis varia species complex. Molecular Phylogenetics and Evolu- tion 120: 103-117. https://doi.org/10.1016/j.ympev.2017.11.014
Zimkus BM, Rédel M-O, Hillers A (2010) Complex patterns of continental speciation: Molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phryno- batrachus). Molecular Phylogenetics and Evolution 55(3): 883-900. https://doi. org/10.1016/j.ympev.2009.12.012
ZooKeys 1201: 167-217 (2024), DOI: 10.3897/zookeys.1201.120750 217